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CHAPTER 6
Cues and decision rules in
animal migration
Silke Bauer, Bart A. Nolet, Jarl Giske, Jason W. Chapman,
Susanne Åkesson, Anders Hedenström, and John M. Fryxell
Ice bars my way to cross
the Yellow River,
Snows from dark skies to climb
the T’ai-hang mountains!
(Hard is the journey,
Hard is the journey,
o many turnings,
And now where am I?)
So when a breeze breaks waves,
bringing fair weather,
I set a cloud for sails,
cross the blue oceans!
Li Po (701–762) ‘Hard is the journey’
Table of contents
6.1
Introduction
69
6.2
Challenges in migration
69
6.2.1
Where to go?
69
6.2.2
When to go?
69
6.2.3
Seasonal and life-cycle migration
70
6.2.4
Travel with or without a predefined target
70
6.2.5
Genetic or cultural transmission of migratory behaviour
70
Cues in the different phases of migration
71
6.3.1
Migration in plankton
71
6.3.2
Migration in insects
72
6.3.3
6.3
Migration in fish
74
6.3.3.1
Life-stage migration in Atlantic salmon
74
6.3.3.2
Feeding migration in pelagic fish: an undefined target
76
Turtle migration
77
6.3.4
68
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6.3.5
Bird migration
78
6.3.6
Migration in mammals
81
6.3.6.1
Migration in bats
81
6.3.6.2
Migration in large herbivores
82
Discussion and Integration
84
6.4
6.1
69
Introduction
The sheer beauty and impressiveness of animal
migrations have long puzzled observers and raised
the questions of how these animals find their way,
what initiates their migrations and how they manage to schedule their journeys at apparently the
right times. There are many challenges that animals
face before and during migration, and they can be
grouped into two major categories namely ‘where to
go?’, dealing with orientation and navigation, and
‘when to go?’, dealing with the timing of activities
and migration schedules. As these questions are
fundamentally different, we also expect different
cues to provide relevant information. Furthermore,
animals make decisions in relation to their physiological state and therefore, cues can be further categorized into internal and external signals.
In this chapter, we tackle these questions not in
terms of why animals migrate (ultimate reasons),
but how they make the right decisions before and
during migration (proximate factors). Migrating
animals rely on external and internal information
such that they can tune their behaviour to their
(changing) requirements and to the development of
their seasonal environments. Here, we show that
these cues, defined as ‘signals or prompts for action’
(Oxford English Dictionary), are not well understood,
although they are most likely highly relevant both
for advancing our fundamental understanding of
migration and for increasing our capacity to manage and conserve migratory systems under the
threat of environmental change.
In the following sections, we first characterize
migration from different perspectives, as different
migration types may require fundamentally different cues and decision rules. Thereafter, we introduce migration in the major migratory taxa that are
readily observable to biologists, i.e. insects, fish,
reptiles, birds and mammals, and seek to identify
the cues that are used for the different steps during
each of their migrations. We finish by highlighting
the general lessons that can be drawn from this
comparative study of cues and decision rules in
migration.
6.2 Challenges in migration
6.2.1 Where to go?
Going along a specific track requires cues for positioning (navigation) and for finding the way
towards the goal (orientation). Important cues for
compass orientation include information from the
magnetic field, the Sun and the related pattern of
sky light polarization and stars, while information
from, for example, landmarks and odours are used
for navigation. Excellent reviews on orientation and
navigation can be found in, for example, Åkesson
and Hedenström (2007), and Newton (2008). However, for many migrating animals we still do not
know how they find their way, which orientation
and navigation abilities they have and which mechanisms they use (e.g. Alerstam 2006; Holland et al.
2006b).
6.2.2 When to go?
Photoperiod has been shown to be involved in the
timing of activities for many species, e.g. initiating ‘Zugunruhe’ (restless behaviour as the premigration phase starts) or determining the speed of
migratory progression. This may come as no surprise, as photoperiod is a reliable indicator of time
of the year and thus can be a useful predictor for the
phenology of resources (Fig. 6.1). Other local and
short-term factors influencing timing of migration include prevailing weather conditions, e.g.
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Latitude
daylight
0 hrs
2
4
6
8
10
12
14
16
18
20
22
24
60° N
40° N
20° N
0°
20° N
40° N
60° N
J
F
M
A
M
J
J
A
S
O
N
D
Figure 6.1 Photoperiod, i.e. day length, as a function of latitude and
time of the year. Day length is here the time between dawn and dusk using
civil twilight (Sun 6° below horizon). The different shades of grey indicate
the day length with darker colours depicting continuous light or permanent
darkness (from Bauchinger and Klaassen 2005).
temperature, wind, drought and precipitation, or
water discharge in rivers, as these factors can significantly influence the costs of the travel ahead
(Chapters 4 and 5).
There are also internal cues that serve as a clock
or time-keeping mechanism. Additionally, physiological state and developmental stage are important
cues as most migrants undergo morphological and
physiological changes in preparation for migration
and internal signals, e.g. hormone levels, indicate
when these changes are completed (Chapter 5).
6.2.3
changes in the body plans often entail a habitat
change and therefore, internal signals are required
that are linked to these developmental processes as
well as cues to locate the next favourable habitat.
However, these organisms may also live in seasonal
environments and, thus, the timing of ontogenetic
processes will also depend on the phenology of the
environment (Skelly and Werner 1990).
Seasonal and life-cycle migration
We can distinguish between two life-cycle patterns
in migratory animals. In one type, which includes
land reptiles, birds and mammals, most body transformations take place within the egg (reptiles and
birds) or within the mother’s womb (mammals),
and the juveniles are only one or a few orders of
magnitude smaller than the adult, and are generally
well suited to the same environment as the adults.
Migration in many of these animals is linked to a
seasonal change in the environment and the cues
involved typically predict these changes.
The alternative pattern includes organisms with
complex life-cycles, such as arthropods, fish, amphibians and sea-reptiles, where animals spawn tiny
eggs that develop into individuals with a body size
several orders of magnitude smaller than the adults,
and with a body form differing substantially from
the adult form. During development, the major
6.2.4 Travel with or without a predefined
target
The best-known type of migration is that between a
few specific localities, e.g. birds between wintering
and breeding sites. In many cases, however, the
migration does not lead to a specific locality or even
to a certain more broadly defined area: In several
species of pelagic fishes, both long-distance feeding
and spawning migrations need not lead to a specific
target. Feeding migration is often driven by continuous local food search (Huse and Giske 1998;
Nøttestad et al. 1999), while the return spawning
migration combines long-distance tracking of preferred spawning sites with physiological constraints
from swimming costs (Huse and Giske 1998; Slotte
and Fiksen 2000). Although some large insect
migrants, such as Lepidoptera (butterflies and
moths) and Odonata (dragonflies) have regular, bidirectional seasonal long-distance migrations that
involve movements that are directed in predictable
ways but not targeted at a specific site or region
(e.g. Chapman et al. 2008a, 2008b; Wikelski et al.
2006), most insect migrations do not even involve
movements in consistent, seasonally preferred
directions.
6.2.5 Genetic or cultural transmission
of migratory behaviour
How do offspring decide where and when to
migrate? Migratory behaviour can be both genetically and culturally determined. In cultural transmission, the young copy their parents’ (or other
group members’) behaviour. Consequently, species
with culturally-transmitted migratory behaviour
are expected to have a social life-style, longer lifespans and (in higher vertebrates) extended parental
care. Prominent examples include schooling fishes
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(e.g. the culturally-induced change of migration
patterns and over-wintering sites in herrings; Huse
et al. 2002), geese and swans among birds (e.g. Von
Essen 1991), and large herd-living mammals such
as antelopes and wildebeest.
Alternatively, migratory behaviour, e.g. routes,
threshold photoperiods, or preferred directions, can
be genetically transmitted when there are no parents, peers, or elders—be it due to high mortality,
short adult life-span, solitary life-style, absence of
parental care or separation of age classes, e.g. age
classes or generations have different requirements
or constraints (differential migration). Examples
of genetic transmission of migratory behaviour
include some birds (e.g. the majority of small passerines and the European Cuckoo Cuculus canorus),
all insect migrants and sea turtles (hatchlings complete their migration alone, and have to return to
the same beach to breed when reaching sexual
maturity many years later).
Besides the general insights, how (part of) migratory behaviour is transmitted is also highly relevant
with regard to global and local environmental
changes. A first review of the effects of environmental change by Sutherland (1998), concentrating on
birds only, showed that none of the species with
culturally determined migration routes had suboptimal routes, i.e. longer than necessary, while
approximately half of the species with genetically
transmitted routes had become sub-optimal. There
is thus a risk that environmental changes may occur
faster than natural selection, particularly for longlived and less fecund life forms. Whether these
findings also apply to other taxa has yet to be
shown. Exceptions to this general pattern appear to
be zooplankton and insects—with their short generation times and high reproductive rates, many
insect pests, for example, are able to adapt to changing conditions rapidly.
6.3 Cues in the different phases of
migration
Migration can be divided into a few major steps—
preparation, departure, on the way, and termination—a cycle that might be repeated if migration is
suspended at intermittent stopover sites. Each of
these steps potentially requires specific cues and
71
decision rules as their demands on the animal’s
physiology and behaviour differ. Similarities might
exist across taxonomic groups in how animals deal
with each of these steps but differences may also be
expected depending on the specific way of migrating or their particular environment.
6.3.1
Migration in plankton
The annual or seasonal migrations in plankton
probably include a higher number of migrants than
any other group (e.g. 1015 individuals of Antarctic
krill Euphausia superba). As an example we present
the much-studied Calanus finmarchius, which is
among the most abundant species of marine calanoid copepods. These North Atlantic copepods
reach an adult body size of a few millimetres, and
spend most of the year in a survival mode in deep
waters. Although the exact depth varies with local
conditions and the state of the individual, and may
range from a few hundred metres to >1000 m
(Kaartvedt 1996), it is vital that they descend deeper
than the winter mixing zone to avoid passive
retransport to the surface layers during winter. The
minimum energetic cost during over-wintering
occurs where the organism is buoyant, so the individual variation in over-wintering depth probably
comes from variation in storage tissue in the form of
wax esters (Heath et al. 2004). Only a small fraction
of the wax esters produced in the preceding feeding
season are consumed during over-wintering (which
is also sometimes called hibernation, diapause, dormancy or resting stage, Hirche 1996). Most is saved
for conversion to eggs in or near surface waters in
spring.
Preparation and departure: These copepods
undergo a series of moults during their life, with
six naupliar stages followed by five copepodid
stages before adulthood. Overwintering is usually
restricted to the fifth copepodid stage (C5). Since
the maximum efficiency in converting food to storage occurs in the C3–C5 stages, the eggs of the overwintering adults must hatch in time to grow and
develop through the naupliar stages in time for
C3–C5 to hit the spring-peak in phytoplankton production. Thus, ascent from deep waters must be
timed well in advance of the peak. Depending on
the food conditions in the surface waters, the
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copepods may produce one or several generations
during spring and summer. Only the last of these
generations will descend to the diapause depth.
This migration pattern is therefore not genetically
hard-wired, but also depends on one or more environmental signals.
On the way and termination: There are still several plausible suggestions for cues involved in the
seasonal migrations for plankton in general and the
species C. finmarchicus in particular. The matter is
further complicated as this latter species lives in a
very diverse range of environments in the North
Atlantic Ocean and adjacent seas, and cues that are
reliable in one area may not be so in another.
Therefore, Hind et al. (2000) modelled the seasonal
dynamics of the species in four different areas: the
North Sea, the Norwegian Sea, the Iceland Shelf
and in a northern Norwegian fjord, to test which set
of cues would produce viable populations in all of
these areas. They found only one set of cues that
produced realistic population dynamics in all areas.
This set consisted of four cues: (i) an external signal;
(ii) an inherited threshold for the downwards
migration; (iii) a physical characteristic of the overwintering depth for the organism (buoyancy); and
(iv) an internal cue for timing of ascent. If ambient
food concentration is above the inherited threshold
value for environmental food concentration, C4
copepodids develop towards adults and another
generation in surface waters. If food levels are
below the threshold, they sink after moulting to C5.
Having reached the over-wintering depth, they
continue to develop at a constant rate, but slower
than for surface dwelling organisms. The cue for
ascent to the surface is that the organism has completed 80% of the development of the C5 stage.
However, one should bear in mind that this is only
an ultimate test (population dynamics modelling)
of the proximate mechanisms—neither the physiological nor developmental mechanisms are understood so far.
6.3.2
Migration in insects
Although migration occurs in all major insect
orders, the actual migrations may often go unnoticed due to the small size of most insects, and the
tendency of many species to migrate at great heights
above the ground. However, the utilization of fast
air currents allows many species to cover enormous
distances (hundreds or even thousands of kilometres), often within just a few days and the consequences of these invisible large-scale insect
movements may be highly conspicuous wherever
they terminate. Some insect migrations are highly
noticeable; among the most impressive of natural
phenomena are the mass migrations in enormous
cohesive swarms of a few species (e.g. the desert
locusts Schistocerca gregaria, the dragonfly Aeshna
bonariensis, and the monarch butterfly Danaus plexippus), which rival the largest flocks and herds of
migratory birds and mammals in terms of biomass,
and far exceed them in total numbers (Holland et al.
2006b).
Insect migrants typically do not make round-trip
journeys, where the same individuals return to their
natal area, nor do most species carry out bi-directional seasonal movements between separate breeding and wintering grounds. Instead, successive
generations engage in windborne displacements
through the landscape, most likely in an attempt to
locate transient and patchily distributed favourable
habitats. The majority of insect migrants take advantage of fast windborne dispersal and fly at altitudes
of from several tens of metres up to a few kilometres above the ground. Relatively few species
migrate predominantly within their flight boundary layer (FBL), i.e. the narrow layer of the atmosphere closest to the ground within which their
airspeed exceeds the wind speed (Taylor 1974)—
this is mostly restricted to large, day-flying species,
such as butterflies and dragonflies (e.g. Dudley and
Srygley 2008).
In most species, migration is restricted to the
adult—winged—life stages and to a single brief
time window of just a few days, due to the short
adult life-span and further because migration typically takes place in the brief period of sexual immaturity immediately following metamorphosis from
the immature stage to the adult (aka oogenesisflight syndrome, Johnson 1969).
Preparation: The development of full-sized wings
and associated musculature is obviously the most
important preparation and many species, e.g.
aphids, have the ability to produce offspring with
varying levels of flight capability in response to
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environmental conditions, e.g. decreasing plant
nutritional quality, and increased crowding. Exceptions to this general pattern exist in longer-lived
species such as the monarch butterfly Danaus plexippus, which builds up substantial fuel reserves by
foraging as adults, and tops these reserves up during intermittent stopover episodes.
The juvenile hormone and its esterase mediate a
range of correlated factors associated with migration, e.g. timing of reproductive maturation, fuel
deposition, development of larger wings and wingmuscles, and increased flight capability.
Departure: Owing to the very short window for
migration in most species, opportunities to choose
the departure time are rather limited and mainly
concern questions of whether to migrate on a particular occasion and at what time of the day to
migrate.
For time of the day, two basic options exist—diurnal migrants take advantage of the higher air temperatures and greater illumination (presumably
facilitating orientation), while nocturnal migrants
benefit from the absence of convective up-draughts
and down-draughts and thus can control their altitude to a much greater extent than day-flying
insects, taking advantage of warm, fast-moving,
unidirectional air currents (Wood et al. 2006;
Chapman et al. 2008b).
The decision whether to initiate migration on any
particular occasion varies between species. Many
insect migrants will not take off when wind speeds
at ground level are too fast (more than a few m/s),
as they cannot control their flight direction immediately after take-off (e.g. green lacewings Chrysoperla
carnea, Chapman et al. 2006). However, as the migration window of most species generally lasts for just
a short period (e.g. two nights in lacewings), they
are unable to migrate if confronted with extended
periods of strong winds, or are forced to do so in
unfavourable conditions.
More complex decision rules are required for species that need to move in a particular direction, e.g.
south in the autumn to escape northern hemisphere
winter conditions. Some species are able to gauge
the presence of favourable high-altitude tailwinds,
facilitating southerly displacement in the autumn.
An example for this is the potato leafhopper
Empoasca fabae—a small insect that is entirely
73
dependent on windborne displacement to escape
deteriorating winter conditions in northern regions
of the US by migrating to its diapause site in the
southern US. Autumn migrants initiate their flights
in response to falling barometric pressure, which is
indicative of the passage of weather fronts that are
followed by persistent northerly air flows, thus
facilitating long-range transport of the leafhoppers
to the south (Shields and Testa 1999).
Green darner dragonflies have a number of simple decision rules that guide their autumn migrations along the eastern seaboard of North America
in a favourable, southerly direction (Wikelski et al.
2006). They initiate migratory flights on days following two preceding nights of dropping temperatures, which are highly likely to be associated with
persistent northerly air flows, and then simply fly
in the downwind direction while avoiding being
carried over large water bodies (and thus out to
sea). Red Admiral butterflies also choose cold northerly tailwinds for their return migrations from
Scandinavia—they fly at high altitudes when fastmoving winds from the north predominate, but low
down when migrating in headwinds (Mikkola
2003).
On the way: Many insect migrants are too slowflying compared with the speed of the air currents
to influence the direction and speed of their movement. The most efficient strategy then is simply to
fly downwind if the migrants are able to perceive
the direction of the current (either through visual
assessment of the direction of movement relative to
the ground, or via some wind-related mechanism).
There is considerable evidence that many high-altitude migrants are capable of aligning their headings in a more-or-less downwind direction (e.g.
Reynolds and Riley 1997), and given that winds
blow in favourable directions, displacement distances will be considerably longer than if the insects
flew across or against the wind (e.g. Wood et al.
2006).
Migrants that fly predominantly within their
flight boundary layer (FBL; Taylor 1974) can control
their direction of movement irrespective of the wind
direction. This is the case for butterflies, which are
powerful fliers and can maintain migration speeds
of 5 or 6 m/s for several hours a day, and for many
consecutive days. To guide their migrations in
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seasonally-favourable directions, these butterflies
must have a compass mechanism. A well-known
example is the monarch, whose eastern North
America population undergoes an annual autumn
migration of up to 3500 km from the late-summer
breeding grounds in eastern Canada and NorthEastern United States to the communal wintering
site in central Mexico. But how do the monarchs orient their flight headings in the correct direction?
Work by Mouritsen and Frost (2002) has demonstrated that autumn-generation monarchs have a
preferred migratory heading towards the southwest, and that during sunny conditions they use a
time-compensated solar compass to select and
maintain this heading. In spring, successive generations of monarchs move progressively northwards
through North America and presumably use the
same orientation mechanism, but in reverse, to
guide their migrations.
Migratory tracks of day-flying FBL insect migrants
(butterflies over the Panama Canal) occur in predictable seasonal directions (from the Atlantic wet
forest to the Pacific dry forest at the onset of the
rainy season), and in at least two species (A. statira
and P. argante) these preferred directions are maintained by reference to a time-compensated solar
compass (Oliveira et al. 1998), i.e. use of visual landmarks on the horizon to compensate for crosswind
drift away from their preferred migration directions
(Srygley and Dudley 2008). Measurements of wind
speed and air speed also indicated that these butterflies adjusted their air speed in relation to wind
speed and their endogenous lipid reserves, so that
they maximized their migratory distance per unit of
fuel (Dudley and Srygley 2008; Srygley and Dudley
2008).
Chapman et al. (2008a, 2008b) have demonstrated
that high-flying migrants, hundreds of metres above
their FBL, can also influence their displacement
direction even though wind speeds far exceed their
own air speed. The moth Autographa gamma is able
to select flight headings that partially compensate
for crosswind drift away from its preferred seasonal
migration directions, thus maximizing the distance
travelled while influencing its migration direction
in a seasonally-advantageous manner. Using a combination of altitude selection to fly in the fastest
winds, and taking up advantageous headings, they
can cover up to 600 km in seasonally adaptive directions during a single night’s flight (Chapman et al.
2008a, 2008b).
Termination: The very act of migration slowly
reduces the inhibition of responsiveness to ‘appetitive’ cues that is typical of migratory flight (Dingle
and Drake 2007), and thus migratory behaviour
itself slowly promotes its own termination. The vast
majority of insect migrants only undertake one, or
at most a few, bouts of migratory flight, and so the
factors that bring about termination of a single bout
of flight are often the same as those that bring about
the termination of the whole migratory phase. These
include depletion of fuel reserves, changes in photoperiod (e.g. nocturnal insects rarely migrate into
daytime, and diurnal species rarely carry on into
night-time: e.g. Chapman et al. 2004; Reynolds et al.
2008), and changes in temperature (e.g. migrations
of nocturnal insects are often terminated due to a
drop in temperature as the night progresses (Wood
et al. 2006)). If the habitat after the termination of the
initial migratory bout is suitable, then that will usually signal the end of the migratory phase, otherwise migration may continue for another bout. In
some species, the flight muscles are autolysed and
converted to increased egg mass after migration, i.e.
they become effectively flightless.
6.3.3
Migration in fish
Although there are about 30 000 species of teleost
fish, only a small fraction of them are currently
known to be migratory. However, these few species
are the dominant marine species in terms of biomass and numbers, and most of the world’s fish
catches are based on them. Many types of migration
exist in fishes, e.g. from freshwater natal areas to the
sea (or vice versa), and between feeding and breeding grounds in the sea. Here we illustrate some
characteristics of fish migration by introducing two
prominent examples, namely life-stage migration in
salmon and feeding migration in herring.
6.3.3.1 Life-stage migration in Atlantic salmon
As predation pressure is considerable in the estuary
and beyond, schooling behaviour is advantageous
and therefore so is size similarity amongst smolt.
This is achieved through growth control by the parr,
Environmental
influence
Genetic
influence
no
Ý
Ý
Ý LBP axis
LBP axis
LBP axis
Circelating
hormones
Circelating
no Ý hormones
Ý
Ý
Circelating
hormones
Endocrine
output
Ion secretory
no Ý protein
mRNAs
Ion secretory
protein
Ý mRNAs
Ý
Ion secretory
protein
mRNAs
Gill gene
expression
NKA activity
&
hypoosencoegulamory
capacity
NKA activity
&
no Ý hypoosencoegulamory
capacity
Ý
Ý
NKA activity
&
hypoosencoegulamory
capacity
Physiological
response
(d)
(c)
(b)
(a)
Figure 6.2 The sequence of events occurring during the parr–smolt transformation (smoltification) in salmon leading to hypo-osmoregulatory development, or not. Here three experimental groups of
Atlantic salmon are presented: anadromous control, parr in February (A) and smolt in May (B) reared under simulated natural photoperiod (SNP); landlocked in May (C) reared under SNP; and
anadromous control in May reared under continuous constant light (D) to demonstrate the importance of brain development of the light–brain–pituitary axis (LBP) early in smoltification on the
downstream endocrine output, gill gene expression and hypo-osmoregulatory capacity. The degree of LBP development is reflected through all downstream processes including physiological development
and Na+, K+-ATPase (NKA) activity (Ebbesson et al. 2007; Nilsen et al. 2007; Stefansson et al. 2007). Reproduced with permisson.
LL
SNP
Landlocked
Anadromous
continuous
light
(LL)
SNP
Anadromous
control
Experimental
groups
Brain pituitary
development
Smolt
Parr
Photoperiod
stimulation
Smolt
Parr
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which is the life stage of young fish in the river
before smoltification. Smoltification is the process
in salmonid parr of preparing for their downstream,
seaward migration and includes a suite of physiological, morphological, biochemical and behavioural changes (Fig. 6.2). Under very benign growth
conditions, smoltification can happen during the
first year of life, but in northern populations this
may take up to seven years. The smoltification decision is based on internal stimuli but the relationship
between the parr’s body condition and the initiation
of smoltification is poorly understood (Stefansson
et al. 2008). According to Thorpe (1977), a bimodality in size appears in the second (or later) autumn.
Only parr larger than 7.5–8.5 cm fork length eventually leave the river as smolt the coming spring. Once
the decision on smoltification is made, the parr
changes into a fast growth mode, where the growth
rate may be 4–5 times higher than before. If the parr
decides to wait, it even goes into anorexia during
the winter (Stefansson et al. 2008).
The next external stimulus is the change in day
length in the following spring, probably combined
with exceeding a temperature threshold. This leads
to growth of the brain and the pituitary gland, leading afterwards to the release of a series of endocrine
hormones. This, in turn, activates gill genes and initiates the physiological processes leading to the
smolt stage (Stefansson et al. 2008). The final downstream migration is triggered by a combination of
light regime, temperature and river discharge (Hoar
1988), leading to simultaneous mass migrations into
the estuary. The smolt will usually remain there for
some months before migrating into the open ocean,
usually as solitary individuals.
How can the salmon find its way back to its native
river, small or large, hundreds of kilometres away
and 1–4 years later? Homing to the river is also
driven by a combination of internal and external
factors. Several hypotheses have been suggested
including a pheromone trail left by out-migrating
fish, counter-current swimming, navigation by
stars, and geomagnetism (Lohmann et al. 2008). It is
quite clear that salmon utilize smell and learned
cues in the later homing phase. Magnetic crystals
have been found in the lateral line sensory system
of salmon (Moore et al. 1990) and in the olfactory
lamellae of trout (Walker et al. 1997), which they
might use for long-distance navigation. Therefore,
they probably use a combination of geomagnetic
information (for long-distance directional migration) and smell and imprinting cues (for choosing
the correct river and stretch of it).
Preparation in parr: While food abundance is the
driving force for the seawards migration in all size
classes, the change from hypo- to hyper-salinity
requires a major transformation of the metabolism
and, additionally, changes in behaviour and skin
pigmentation as the young salmon transforms from
a bottom-dwelling territorial parr into an openwater schooling smolt. As this decision is taken long
before the actual migration, cues for preparations
come from both internal and external sources—once
a threshold body condition (size) is reached, daylength initiates the onset of body and metabolism
changes.
Departure in smolt: After having completed all
body changes, the smolt often waits for the autumn
river discharge to depart and go downriver
seawards.
On the way smolt: Seawards, the smolt remains
for some time at the estuary to become imprinted
and then leave for the ocean in groups, where they
become solitary again and mainly follow food. On
their way back, they probably initially use some
sort of magnetic field orientation, and gradually
change to olfactorial orientation when they are near
the home-river (e.g. Healey and Groot 1987).
Termination in returning adults: Once they have
arrived in their target area in the natal river, migration is suspended.
6.3.3.2 Feeding migration in pelagic fish: an
undefined target
In several species of pelagic fish, long-distance
migrations may be directional rather than to a specific localizable target, e.g. mackerel Scomber scombrus
and blue whiting Micromesistius poutassou migrate
northwards from spawning areas around the British
Isles into feeding areas in the Norwegian Sea.
Thereby, they benefit both from the later spring and
summer further north, and also from the gradual
increase in daylight-hours in the northern summer.
Both factors contribute to prolonged high feeding
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and growth rates (Nøttestad et al. 1999). Similarly,
herring Clupea harengus migrate westwards from the
mild Atlantic waters off the coast of Norway towards
the colder waters in the west, with delayed spring
production (Varpe et al. 2005). For some decades, the
whole adult population of Norwegian spring spawning herring has been over-wintering in the deep
Tysfjord in northern Norway. Spawning migration in
spring is southwards along the coast of Norway. The
further south the eggs are spawned, the better the
prospects for larval growth and survival. However,
as migration is energetically costly, there is a tradeoff between fecundity and migration distance such
that small individuals migrate shorter distances and
larger individuals longer, i.e. further south (Slotte
and Fiksen 2000). Hence, the likely cues are a combination of physiological state (spawning migration in
herring) and a seasonal signal such as day length
(end of northwards or westwards feeding
migration).
Very little, if anything, is know about how fish
find their way and make decisions. Geomagnetism
has been proposed for long-distance navigation
(e.g. Lohmann et al. 2008). Many of the species also
migrate in large schools, which may act as cooperative units for food searching (Clark and Mangel
1986) and decision-making (Huse et al. 2002). Thus
schooling acts both to reduce predation risk, and
to increase the chance of being on the right track
for future food resources. During the feeding
migration, models indicate that a long-distance
direction finder may not be needed as the fish simply follow the seasonal development of the food,
which will automatically lead them to profitable
places (Huse and Giske 1998). However, models
also indicate that a separate ‘homing motive’ is
needed for the return migration, during which
local gradients in food or temperature may not be
helpful (Huse and Giske 1998). Unfortunately, it is
not known whether the decision to return is based
on some seasonal signal or the state of the organism, or both.
6.3.4 Turtle migration
Long-lived sea turtles regularly commute between
two completely different environments, the open
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ocean for foraging and sandy shores for egg-laying. Some sea turtles, e.g. the leatherback turtle
(Dermochelys coriacea), spend several years foraging in pelagic habitat (e.g. Hays et al. 2004) and
accumulate body stores, which they later use for
mating and producing eggs. These two habitats
are usually separated by vast areas of unsuitable
habitat and, consequently, migrations are often
long.
Others, e.g. herbivorous green turtles (Chelonia
mydas), also lay eggs on sandy beaches but forage
as adults on sea grass and algae along shallow
coastal areas (e.g. Mortimer and Carr 1987, Bjorndal
1997). One of the longest distance migrations is
performed by Ascension Island green turtles,
migrating between breeding sites at Ascension
Island and foraging areas along the Brazilian and
Uruguayan coasts (Carr 1984; Papi et al. 2000;
Luschi et al. 2001).
It is well known that sensory information is important for navigation by hatchling sea turtles when
they depart to the sea (Lohmann and Lohmann 1996;
Lohmann et al. 2008), but it is less well understood
what information is used by the adults when returning to breed (Luschi et al. 2001; Åkesson et al. 2003).
Even less is known about the migratory behaviour
and information used by sub-adult sea turtles
(Godley et al. 2003) and how the transition takes
place from the genetically programmed guidance of
the hatchlings into the migration programme guiding the sub-adults and adults later in life (Åkesson et
al. 2003). Most likely, the turtles use partly genetically
encoded behaviours, but also learn to incorporate a
number of cues into their navigational toolbox
(Åkesson et al. 2003).
Preparation: Many sea turtles need several years
to recover from a major migration and egg-laying
event and during this time they store fat as fuel. For
example, female Ascension green turtles migrate to
the island to lay eggs, where they do not forage at
all for 5–6 months. Apparently they exhaust most of
their reserves during the event, such that their
recovery and preparation for the next migration
and breeding bout requires approximately 3–4 years
(Carr 1984).
Departure: Hatchlings: When the hatchlings in a
clutch escape from the nest, they first climb to the
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surface of the sand during the day and await the
night. At that stage they are stimulated by their
nestmates’ movements such that all siblings leave
at night in a synchronized fashion. Once in the
water, they mix with other hatchlings and depart on
their independent migratory journeys. When they
depart to open sea, their movement is both active
and passive, i.e. partly swimming and partly drifting with the currents. The timing of departure relative to the season very much depends on the timing
of egg-laying, which again depends on the foraging
conditions encountered in the wintering areas
(Godley et al. 2001).
On the way: Sea turtles have been shown to use a
number of different cues to orientate and navigate
during migration (e.g. Lohmann and Lohmann
1996, Åkesson et al. 2003). Loggerhead turtle hatchlings (Caretta caretta) respond to light when leaving
the sand and moving along the beach; later they
have been shown to swim against the waves to
leave the shore and, once they are in more open
water, they probably use magnetic field information
for navigation as has been shown in experiments
manipulating the magnetic field (Lohmann et al.
1999, 2001). Studies on adult green turtles have tried
to identify cues used during migration, but also
when searching for the breeding island after displacement. It was found that successfully homing
turtles responded to local information, suggesting
they are using information carried with the wind
from the island (Papi et al. 2000; Luschi et al. 2001;
Åkesson et al. 2003).
Termination: For all sea turtles, breeding migration ends as soon as they have reached the breeding
grounds. During foraging migrations, differences
exist between the pelagic species, such as the leatherback turtle, that can be considered to be constantly
moving and exploring the open ocean environment
(Hays et al. 2004), and coastal foragers, such as the
green turtle, which forage along shallow coastal
sea-grass beds.
6.3.5
Bird migration
The classic bird migration is the biannual migration
between breeding and wintering grounds. The
breeding grounds are suitable for nesting and hatchling/fledgling survival, whereas the wintering
grounds are more suitable for post-fledgling and
adult survival. Because birds are able to fly, they can
travel long distances relatively cheaply and quickly
(Chapter 4), e.g. the longest non-stop migratory
flight recorded is that by bar-tailed godwits (Limosa
lapponica) crossing the Pacific Ocean from Alaska to
New Zealand, a distance of more than 10 000 km
(Gill et al. 2009).
Exceptions to this are moult and facultative
migrations. In moult migrations, birds appear to
migrate to predator-free areas where they can safely
shed their flight feathers. In facultative migrations,
birds only migrate long distances when food is
sparse, e.g. many finches (Newton 2006). At an
extreme end of the spectrum are birds that are
nomadic, like the grey teal (Anas gracilis) looking for
ephemeral water and food sources in a desert landscape in Australia (Roshier et al. 2008).
Two main flight modes exist—flapping and soaring—each having particular consequences: Flapping
flight is very costly but can be used under a wide
range of weather and topographic conditions,
whereas for soaring, thermals or wind are needed
(Chapter 4).
The majority of birds cannot feed while flying,
and in many cases the total travel distance exceeds
the maximum flight distance. Thus, the birds need
stopover sites where they can replenish their
reserves. A good example is tundra swans (Cygnus
columbianus), which migrate 4000–5500 km (Nolet
2006), whereas their maximum recorded non-stop
flight is 2850 km (Petrie and Wilcox 2003). These
swans mainly refuel on energy-rich, below-ground
parts of macrophytes in shallow lakes and wetlands
along the route (Beekman et al. 1991).
Preparation: Before actually embarking on migration, most birds partly change the composition of
their bodies, e.g. increase flight muscles at the
expense of leg muscles, atrophy digestive and metabolic organs (Piersma and Gill 1998; Biebach 1998;
van Gils et al. 2008; Bauchinger and McWilliams
2009) and accumulate body stores. Photoperiod is an
important external signal for preparations; it has
been shown to initiate ‘Zugunruhe’, i.e. migratory
restlessness in many migratory passerines (Gwinner
1990), but also many geese, swans and waders start
accumulating body stores, altering their digestive
system and building up flight muscles from a
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particular day length onwards. The specific value of
day length at which these transformations are
started is under strong genetic control, as evidenced
by hybridization, parent–offspring comparisons and
effects of changing selection pressures (Newton
2008). Birds kept under constant day length for up to
several years still showed a circannual rhythm with
the right sequence of annual events (migratory fat
deposition and restlessness, gonad development,
and moult) suggesting that getting into the migratory state is under internal control (Gwinner 1977).
But these cycles tend to drift and be either shorter or
(most often) longer than a calendar year. This internal control is most rigid in long-distance migrants
that are normally confronted with most variation in
day length.
Thus, under natural conditions the exact timing
of events is most likely determined by a combination of internal and external factors such that the
internal system is adjusted by seasonal changes in
photoperiod, as has been shown with experiments
with extra light or shorter than annual cycles
(Newton 2008).
Departure: The exact timing of migratory departure is fine-tuned by secondary factors like temperature, wind, rain and food supplies (Newton 2008).
Birds have been shown to choose favourable flight
conditions and preferably leave on days with tailwinds and no rain. In the Swainson’s thrush (Catharus
ustulatus), departure decisions are best predicted by
both a high daily temperature (>20°C) and low wind
speeds (<10 km/h) at the time of presumed take-off.
If one of these conditions is not met, the individual
will not take off. However, such apparently strict
rules also lead to serious errors, e.g. individuals take
off at low local winds, and yet ascend into air streams
that will push them backwards against their flight
direction (Cochran and Wikelski 2005).
One means by which birds may forecast improving weather conditions before they actually occur
has been hypothesized to be sensing air pressure
changes (Newton 2008, Keeton 1980). In facultative migrants, departure may also be delayed until
weather conditions for refuelling deteriorate (Newton
2008; Gilyazov and Sparks 2002).
The decision to depart from a stopover site is
probably based on rather simple behavioural rules.
Passerines that lose or increase fuel stores at a high
79
rate leave a site quickly, whereas the intermediate
birds stage the longest (Schaub et al. 2008). Geese use
a mixture of endogenous and external cues, with the
endogenous cues having a stronger effect as the season progresses (Bauer et al. 2008, Duriez et al. 2009).
On the way: Birds have been shown to use several
cues to guide them in the right direction on longdistance migrations. The combination of cues may
be essential for correct navigation as directional
cues change with place (e.g. the magnetic compass;
Wiltschko and Wiltschko 1972) and time (e.g. the
sun compass; Kramer 1959). Birds use a combination of cues for recalibration. For instance, recent
experiments suggest that birds use cues from the
setting Sun to re-calibrate their magnetic compass
before migrating at night (Cochran et al. 2004). At
sunrise or sunset, birds can use skylight polarization, especially visible close to the horizon, as a
compass (Able 1982; Muheim et al. 2006). Uniquely
to birds, star patterns that indicate the axis of rotation of the night sky have been hypothesized to be a
directional tool (Newton 2008). However, laboratory-based experiments on stellar orientation in birds
could also be explained by the fact that individuals
have the rule to use the single brightest and nonmoving light as the main orientation cue.
If birds do not compensate for the change in local
time when travelling across longitudes, a sun compass would lead them along routes similar to great
circle routes (Alerstam and Pettersson 1991). In contrast, if they compensate and reset their internal
clocks regularly while crossing longitudes, they
would follow a constant rhumbline route (Fig. 6.3).
Birds following one (Alerstam et al. 2001) or the
other (Green et al. 2002) have been found.
The direction of migration is also under endogenous control (Gwinner and Wiltschko 1980; Helbig
1991). The direction is reversed when the return
migration starts (Newton 2008). Together, the inherent period and direction of migration result in naive
migrants being able to stop in the right area. Juvenile
starlings that were trapped on migration and displaced by aeroplane to Switzerland, east of the
usual wintering area of the population, continued
their migration in the same direction and over the
same distance that they would otherwise have
flown and ended up in southern France or Spain
(Perdeck 1967). In contrast, the trapped adult star-
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Figure 6.3 Two extremes: a global view of the Earth from space (insert; orthographic projection centred in the Wadden Sea 54 °N, 8.5 °E), and a
Mercator projection flattening and stretching the globe. On the Mercator projection all straight lines are constant geographic bearings (loxodromes,
rhumblines), whereas straight lines through the centre of the orthographic projection represent great circles (orthodromes). Note that the scale at the
Equator is similar on both maps (from Gudmundsson and Alerstam 1998). Great circle routes are thus the shortest distance between any two points on the
Earth’s surface whereas rhumbline routes may be easier for navigation as they require no re-adjustments of headings but always cross meridians at the
same angle. The difference in distance between both routes is small (<1%) at high latitudes and distances of less than 30° longitude but increases
significantly thereafter. For instance, for travelling along 50° latitude and across 180° longitude, the rhumbline route is 45% longer than a great circle route.
lings were found in their traditional wintering area
in northern France and England, so they must have
used goal orientation. Interestingly, the juveniles
returned to their new wintering area in the subsequent years, showing they also switched to goal
orientation in later life. Similarly, white-crowned
sparrows (Zonotrichia sp.) were caught in Washington
while migrating from Alaska to California, and
were flown to the US East coast. From there, adults
headed back towards Californian wintering
grounds, whereas juveniles headed south, presumably in an innate direction (Thorup et al. 2007).
Some migrations require changes in direction or
migratory steps along the way, e.g. to avoid inhospitable environments. Birds from populations that
change direction during migration show a corresponding change in direction in orientation cages as
the season progresses, indicating that this is also
under genetic control (e.g. Gwinner 1977; Helbig
1991). In other cases, local conditions serve as cues.
For instance, pied flycatchers Ficedula hypoleuca
changed their directional preference only when
confronted with the magnetic conditions where
they normally change direction, and not when mag-
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Figure 6.4 Adult dark-bellied Brent geese (Branta b. bernicla) arriving at their still largely ice-covered breeding grounds in Taimyr, northern Siberia. In
experienced birds, the decision to stop migrating is influenced by cues indicating that a familiar nesting locality has been reached (Photo: Andries Datema,
Alterra Wageningen-UR).
netic conditions were kept constant (Wiltschko and
Wiltschko 2003). Thrush nightingales Luscinia luscinia that were captured in southern Sweden at the
start of their first autumn migration were exposed
either to the local magnetic field or to an artificial
magnetic field typical of northern Egypt, where
they are thought to prepare for crossing the Sahara
(Fransson et al. 2001). The latter group responded
by accelerating fat deposition, suggesting that there
is a built-in genetic response to local conditions.
Replenishment of the fat store itself may act as a
cue to continue migration: in several experiments it
was demonstrated that migratory restlessness and
inclination to leave were higher in fatter than leaner
individuals. Most of these studies were performed
at localities where the birds were preparing for a
major crossing (Newton 2008).
Termination: When tested under identical conditions in the lab, the duration of migratory restlessness is longer in long- than in short-distance
migrants, even within species (Berthold and Querner
1981). Cross-breeding experiments showed that this
is an inherited trait (see also Berthold 1999 for an
experiment with hybrids of redstarts). In birds from
the same species, those wintering furthest away
from the breeding grounds show a tendency to start
spring migration earlier (King and Mewaldt 1981).
Juvenile blue-winged teal Anas discors caught in the
autumn and held captive for a while, migrated less
far than normal after release at the same site
(Bellrose 1958). This shows that the decision to stop
is at least partly under genetic control. However, in
adult birds the opposite was found, with the migratory restlessness continuing longer than normal
when held captive for a while during spring migration (Newton 2008). Also, when held at the breeding location, indigo buntings Passerina cyanea did
not migrate after release in the spring, whereas the
control birds that were displaced 1000 km to the
south did (Sniegowski et al. 1988). The same was
true for white storks Ciconia ciconia reared in captivity and released in a reintroduction programme
(Fiedler 2003). In experienced birds, the decision to
stop is therefore apparently influenced by cues indicating that the familiar locality has been reached
(Fig. 6.4).
6.3.6
Migration in mammals
6.3.6.1 Migration in bats
Even though bats are mammals, their ability to fly
makes them more like birds in terms of their opportunities for and ecology of migration, but relatively
little is known about their migration biology in
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comparison with birds. This is probably because
temperate bats have adopted hibernation as their
main strategy for surviving periods of resource
depression in a seasonal environment. Yet, in the
family Vespertilionidae, migration occurs in 23 out
of 316 species classified (7%), and has evolved in 15
genera with apparently little phylogenetic inertia
(Bisson et al. 2009). The distances of bat migrations
are shorter than for birds, with maximum migration
distances around 2–3000 km (Hutterer et al. 2005).
In temperate bats, long-distance migration occurs
mainly in species that use trees for roost sites
(Fleming and Eby 2003), but in these species migration is combined with hibernation at the wintering
site. Migration also occurs in tropical species but
movement distances are generally rather short and,
in most cases, are driven by the phenology of fruiting trees (Fleming and Eby 2003). Differential migration is common in bats, with females migrating
further north than males to raise their young, presumed to be due to higher resource needs for raising the young (Fleming and Eby 2003). As in birds,
partial migration also occurs, i.e. part of the population migrates and the other part is resident.
Preparation. Bats accumulate fat deposits before
hibernation (e.g. Kunz et al. 1998), and therefore fat
is probably the main fuel used during migration
(McGuire and Guglielmo 2009).
Departure. Bats most likely depart during the
early night hours, similarly to nocturnally migrating birds. Departure conditions are little studied,
but is seems as if migration activity is highest when
wind speeds are low (Petersons 2004).
On the way. It has recently been shown that bats
possess a magnetic sense (Holland et al. 2006a), and
it therefore seems likely that this is involved in orientation during migration. Otherwise, next to nothing is known about orientation and navigation in
bats, although recent evidence suggests that the
greater mouse-eared bat Myotis myotis calibrates a
magnetic compass with sunset cues (Holland et al.
2010). Some frugivorous species seem to track the
phenology of their main food source on migration
(Fleming and Eby 2003).
Long-distance migration generally consists of
several cycles of fuelling followed by migratory
flight. It remains to be shown whether bats follow
this model, but there are some indications that they
do stop over for fuelling (Petersons 2004). Since bats
are mainly nocturnal they must divide their active
period (the night) between foraging and migratory
flight during migration. Because migrating bats’
rate of energy consumption is typically much higher
than that of fuel accumulation, it is expected that
the proportion of time spent on stopovers should be
much longer than that spent on migratory flights
(Hedenström 2009). One way of saving energy on
migration is to use torpor during periods of fuelling, i.e. lowering the body temperature during
daytime roosting and thereby increasing the net
rate of fuel accumulation (and hence overall migration speed).
In flight, there are alternative ‘optimal’ flight
speeds predicted from flight mechanical theory
(Hedenström 2009), with the maximum range speed
being the best option for minimizing energy cost
per unit distance. A comparison between foraging
and commuting flights in Pipistrellus kuhlii showed
that these bats select flight speed according to this
prediction. Overall, bats seem to fly at slower speeds
than birds of similar sizes (Hedenström et al. 2009).
The overall migration speed includes time for
fuelling and flight (Hedenström et al. 2009), and is
predicted to be about 46 km/day on the basis of fuel
accumulation rate, energy consumption during
flight and flight speed. Ringing recoveries of
Nathusius’s bat P. nathusii showed a migration
speed of 47 km/day (Petersons 2004), which is comparable to that of short–medium distance migrating
birds. Flight altitudes of bats on migration appear
to be rather low (Ahlén et al. 2009), although freetailed bats Tadarida brasiliensis may reach altitudes
of about 3000 m when foraging (Williams et al.
1973).
Termination. Birds have an inherited migration
programme that determines when to cease migration, but whether bats have a similar mechanism is
not known.
6.3.6.2 Migration in large herbivores
Seasonal nomadism and migration have been documented numerous times in terrestrial mammalian
herbivores and occur on every continent (Fryxell
and Sinclair 1986). There are three types of situation
in which herbivore nomadism or migration are common, perhaps even typical. The first situation is
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species inhabiting montane environments, such as
elk, mule deer; red deer, and montane ecotype caribou (Albon and Langvatn 1992; Brown 1992; Horne
et al. 2007; Hebblewhite et al. 2008). Migration is
relatively common in herbivore species inhabiting
open savannah or tundra environments, such as
tundra ecotype caribou in North America, wildebeest, zebra and Thomson’s gazelles in the Serengeti
and Tarangire ecosystems of Tanzania, white-eared
kob and tiang in the Boma ecosystem of Sudan, and
Mongolian gazelles (Pennycuick 1975; Inglis 1976;
Fryxell and Sinclair 1986; Durant et al. 1988; Fryxell
et al. 2004; Boone et al. 2006; Mueller et al. 2007;
Holdo et al. 2009). Finally, seasonal migration by
ungulates also occurs in temperate regions subject
to severe climatic variability, such as woodland caribou, pronghorn antelope, saiga antelope, whitetailed deer or mule deer (Rautenstrauch and
Krausman 1989; Nelson 1998; Johnson et al. 2002;
Ferguson and Elkie 2004; Berger et al. 2006; Sawyer
et al. 2009; Singh et al. 2010).
Seasonal onset of vegetation growth is strongly
temperature-dependent in montane ecosystems in
temperate to arctic regions. As a consequence, snow
melt occurs later at high elevations and vegetation
growth is retarded relative to lower elevations
(Pettorelli et al. 2005). It is common for terrestrial
herbivores to exhibit seasonal shifts in accordance
with seasonal green-up (Albon and Langvatn 1992;
Horne et al. 2007; Hebblewhite et al. 2008; Berger
et al. 2006; Sawyer et al. 2009). In low-lying forbs and
grasses, structural compounds, such as lignin and
cellulose, are incorporated more and more into stem
and leaf tissues as the plant grows taller, reducing
digestibility and lengthening the processing time in
herbivore digestive tracts (van Soest 1982). As a
consequence, optimal rates of nutrient intake can
often be best achieved by feeding on relatively
immature ramets (McNaughton 1984; Hobbs and
Swift 1988; Fryxell and Sinclair 1988; Illius and
Gordon 1992). By appropriate timing of migration
up the elevation gradient, herbivores are able to
access young vegetation and maintain optimal
nutrient intake over a prolonged period. Over the
course of the winter, animals usually retreat to lowlying areas, which are less exposed to severe climatic conditions and have residual plant standing
crop from the growing season. A similar pattern is
83
seen in tundra systems, with animals retreating to
woodland margins during winter, but venturing far
out on to the tundra during the brief growing
season.
In savannah environments, animals usually follow rainfall gradients, from the more arid rangelands used during the brief growing season to
higher rainfall areas used during the driest part of
the year (Pennycuick 1975; Fryxell and Sinclair 1988;
Mueller et al. 2007). As nutrient quality is often
inversely related to annual rainfall levels (Bremen
1983), migrants are able to access young vegetation
at an optimal growth stage during the growing season in the arid areas, while retreating to high rainfall areas when arid lands dry out. Nomadism is
characteristic when rainfall is unpredictable in
space (Fryxell et al. 2004; Mueller et al. 2007, 2008),
though this is often superimposed on a relatively
dependable migratory pattern at coarser temporal
and spatial scales (Wilmshurst et al. 1999; Boone
et al. 2006; Holdo et al. 2009).
In temperate regions with extreme seasonal variation in climate, it is common to see migration from
summer home ranges, presumably chosen primarily to obtain food and reduce predation risk, to
winter ranges with less snow cover or improved
shelter from snow and wind (Rautenstrauch and
Krausman 1989; Nelson 1998; Johnson et al. 2002;
Ferguson and Elkie 2004; Berger et al. 2006; Sawyer
et al. 2009, Singh et al. 2010).
It seems likely that there is at least some learned
or cultural component to migration behaviour in
terrestrial herbivores, though this question has
received relatively little attention in the ungulate
literature. The cultural conjecture is based on welldocumented examples of altered migration routes,
adoption of migration by previously resident animals and even cessation of migration within a single generation (Nelson 1998). Longitudinal studies
clearly suggest that partial migration is typical of
northern white-tailed deer, with young individuals
typically mimicking the migratory behaviour of
their mothers, but capable of shifting to different
strategies (resident or mixed) later in life (Nelson
1998). Similarly, elk in Banff National Park were
largely migratory before the 1990s (Woods 1991).
Re-invasion of wolves into areas in the Bow Valley
from which they had been extirpated led to a
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dramatic change in space use patterns by elk over
the course of 10 years (Hebblewhite et al. 2005), with
most individuals concentrating year-round near
towns that provided security from predation as well
as improved nutrient intake on a year-round basis
(McKenzie 2001; Hebblewhite et al. 2005, 2008).
Preparation and departure. Because large herbivores are highly mobile, feed while they travel, and
have relatively slight energetic costs of movement
relative to other taxa, there is little indication of
extensive physiological preparation for seasonal
movements. Cues for the initiation of migration and
nomadism are thought to include seasonal changes
in temperature, precipitation, and water quality
(Pennycuick 1975; Rautenstrauch and Krausman
1989; Albon and Langvatn 1992; Nelson 1998;
Wolanksi and Gereta 2001; Mahoney and Schaefer
2002; Boone et al. 2006; Gereta et al. 2009), although
evidence is largely anecdotal. For example, Serengeti
wildebeest have been seen to reverse direction and
return to previously vacated areas when temporary
periods of drought interrupt the usual onset of the
rainy season (Pennycuick 1975). This suggests that
rainfall is a key variable for this species, but it is
hard to disentangle that from other putative causal
variables (young vegetation abundance, water quality) that co-vary with rainfall.
On the way. Once underway, it is not clear what
proximate cues migratory herbivores use to guide
their movements. Some species have specific migration routes that are travelled year after year, such as
pronghorn antelope in the mountains of Wyoming,
Idaho, and Montana (Berger 2004, 2006), mule deer
in Wyoming (Sawyer et al. 2009), and montane caribou in Alaska (Horne et al. 2007). In each case, specific individuals travel the same corridors as they
shift from winter to summer ranges. Anthropogenic
habitat changes that create bottlenecks in such
migration corridors are a source of considerable
conservation concern, because there are clear examples of migrants being negatively affected by anthropogenic barriers to movement (Williamson et al.
1988; Mahoney and Schaefer 2002; Berger et al. 2006;
Ito et al. 2005, Chapter 11).
Migration routes in other systems seem much
less repeatable within individuals from year to year
(Wilmshurst et al. 1999; Thirgood et al. 2004; Boone
et al. 2006), suggesting cues may be regional in
nature. For example, movement trajectories of
Serengeti wildebeest and Thomson’s gazelles can
be fairly precisely predicted in coupled map lattice
models on the basis of local rainfall, grass biomass
and soil nutrient levels (Fryxell et al. 2004; Holdo
et al. 2009), but only when animals are capable of
choosing new locations to move to within ranges of
the order of 100s of km2. Smaller zones of perception would probably lead individuals to concentrate
in areas of local fitness peaks, thereby disrupting
the migration that is repeatedly observed at a
coarser spatial scale. In a particular area, residents
can prefer different habitats from migrants (M.
Hebblewhite, pers. comm.), suggesting either that
migrants and non-migrants differ in their selective
constraints or that migrants are unable to choose
the best local resources because of lack of familiarity with the area.
6.4 Discussion and Integration
In this chapter, we have considered the cues that are
used in several phases of migration across taxonomic groups. Although naturally many differences
appear due to the specifics of each species’ migration, considerable similarities appear to exist in the
cues involved in the different phases of migration
(Table 6.1).
In all species, preparations for migration involve
entrainment to time of the year, as all environments
are seasonal to some degree, thus particular times
are more suitable for particular activities. Indeed,
even at very low levels of seasonality, animals
should migrate in order to make use of the varying
levels of food in different areas (Barta et al. 2008).
Therefore, the occurrence of photoperiod as a cue in
almost all taxa is not surprising.
However, as migration is a daunting activity in
the life-cycle or annual cycle, it also requires bodily changes, such as the accumulation of energy
stores, the build-up of the locomotion apparatus—
often at the expense of the digestive and/or reproductive system, and the transformation of a
freshwater- to a seawater-adapted life-form or the
achievement of a particular developmental stage.
Whenever these changes are accomplished, an
internal cue is produced indicating that the animal
is ready to depart.
Photoperiod, Crowding during immature
stages, Habitat deterioration (food,
predation or parasitism)
Reach minimum body size; for some:
physiological adaptation to new
environment
Light regime, internal status, and
migratory restlessness
Photoperiod, Build-up flight apparatus,
Reduction digestive system
Insects
Fish
Large mammals
Mammals: Bats
Birds
Salmon: Autumn river discharge
Descending migration: Food level below
threshold
Ascending migration: C5 stage to 80%
developed
Favourable flying conditions, e.g. tailwinds
Departure
Local food search, or long-distance
spawning location tracking
Time-compensated sun compass
Continue until buoyancy depth and below
mixing depth
On the way
Migration reduces inhibition to appetitive
cues (in mig. bout); depletion of fuel
reserves, changes in photoperiod or
temperature
Arrival in locations favourable for spawning
Descend: arrival in buoyancy depth; Ascend:
arrival in surface waters
Termination
Favourable departure conditions, e.g.
Visual information (bright skylight),
Arrival on specific target location, e.g.
night, with currents
direction of waves, geomagnetic field, wind feeding or wintering grounds
Favourable flight conditions (wind, rain,
Sun compass, magnetic field, skylight
Arrival on specific target location, e.g.
air pressure). Fuelling rate and body stores. polarization, star pattern. Direction
breeding or wintering grounds. Naïve birds
Cumulative temperature or
under hormonal control, sometimes
have an inherent migratory period
related proxy
responses to local conditions
Accumulate fat deposits (torpor during
Early night hours, low wind speeds
Not much known, probably use magnetic Unknown
fuelling periods)
field calibrated by direction of sunset
No particular (physiological) preparations Seasonal changes in temperature,
Not much known; may follow gradients.
Unknown
precipitation and water quality but
evidence anecdotal
Undergo all naupliar stages and reach
5th copepodid stage.
Storage of wax esters
Plankton: Calanus
finmarchicus
Turtles
Preparation
Cues for
Table 6.1 Summary of the cues identified for the four major steps of migration, in all major migratory taxa
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For the actual departure, another external cue is
often involved, which is usually related to travel
conditions, e.g. wind, precipitation, temperature.
Thus, animals prefer to depart during periods of
favourable conditions, for instance, flying animals
wait for tailwinds in their preferred directions; swimming animals use river discharge or sea-currents.
On the way, orientation and navigation determine the migration route taken but they may also
be involved in indicating when migration is to be
terminated. Animals heading for a specific location
need to recognize this location, which is an option
only for experienced animals, whereas naive individuals (e.g. first-time migrants) need to have a
genetic programme that signals when to stop.
Alternatively, migrations without clear endpoints,
e.g. between feeding locations, may involve physiological cues for the termination of migration. Here
again, internal signals play a greater role as they
indicate when a threshold state is reached, e.g. sufficient body reserves have been accumulated for a
subsequent breeding attempt.
Although we can make very rough generalizations such as these, we need to realize that currently
we know the full set of cues and decision rules used
throughout their annual cycle for hardly any species. For most species, we don’t have any idea which
cues and decision rules, orientation and navigation
mechanisms they use during (specific parts of) their
migration. However, such knowledge is all the more
urgently required in the face of human-induced
environmental changes. These changes affect the
size and quality of habitats as well as the distances
that separate suitable environments. Furthermore,
climatic conditions are changing, but to complicate
matters some areas on the globe are expected to be
affected much more than others. For migratory animals, such changes pose particular challenges as
they visit multiple, distant sites during their annual
or life cycles—often even in different ecosystems. If
we are to predict the consequences of such changes
for migratory animals, we need to close the gaps in
our current knowledge and gain a thorough understanding of the cues and decision rules used during
migration as well as animals’ orientation and navigation mechanisms.
To this end, we need to overcome the considerable bias both in the species and taxa studied and
also in the type of questions asked and the
approaches used. Birds are by far the best-studied
taxon at present, followed by the economically relevant fish species, while comparatively little is
known for the other taxa.
Most studies on navigation, orientation and decision rules have so far been conducted in captivity.
Although such studies can provide important first
indicators of the processes involved in natural
migration, the relative importance of different cues
can only be established in complex environments.
Hence, it will be essential to study migratory decisions of wild, naturally migrating individuals
(Wikelski et al. 2007).
Traditionally, migrations of animals (in particular, birds) have been identified using recoveries and
resightings of marked individuals. Especially for
larger birds such as swans and geese, individual
marking, e.g. with neck-rings, has been possible,
allowing detailed observations along their routes.
More recently, the advent of increased communication possibilities and technological advances have
led to significant progress in, for example, the development of satellite transmitters, geolocators, or the
miniaturization of existing devices such that the
movement of individuals of smaller and/or clandestine species can be followed in great detail (e.g.
discovery of migratory routes in turtles, Hays
2008).
Data obtained with these devices can provide
insights into the individual level of decision-making
involved in the different steps during migration
and thus provide mechanistic rather than phenomenological insights (Chapter 8). Such individual
movement data can be analysed across taxa (www.
movebank.org). Furthermore, these tracks can be
integrated with detailed geographical and dynamic
meteorological information allowing the identification of both the internal and external (environmental) determinants of migration decisions.
Another avenue for further advances in our
understanding is improved integration of theoretical and empirical efforts (Bauer et al. 2009;
Chapter 8). Significant progress in science has
often been achieved when theoretical developments have inspired new experiments or when
startling empirical findings have inspired the
development of new theories. Despite pioneering
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D I S C U S S I O N A N D I N T E G R AT I O N
efforts (e.g. Alerstam and Lindström 1990), the
interaction between theoreticians and empiricists
has been too limited to date in the study of animal
migration. Several modelling approaches exist,
ranging from simple optimality models (e.g.
Alerstam and Hedenström 1998), dynamic optimisation models (e.g. Houston and McNamara
1999), game-theoretic models (e.g. Kokko 1999),
individual-based models (e.g. Pettifor et al. 2000)
to models based on evolutionary methods (genetic
algorithms and neural network models, e.g. Huse
et al. 1999). Again, the use of these models has
been highly biased, with birds being the most
studied taxon with the widest variety of theoretical approaches used.
87
Methods for the identification of cues and decision
rules are numerous and include (but are not restricted
to) translocation/displacement experiments (e.g.
Luschi et al. 2001), cross-breeding experiments (Helbig
1991) and a combination of theoretical and empirical
approaches (confronting models with data), e.g. simulation models (e.g. Duriez et al. 2009) and proportional hazards models (e.g. Bauer et al. 2008).
We believe that much could be learned by overcoming taxonomic borders and integrating theoretical and empirical efforts—particularly in our
rapidly changing world that challenges migratory
animals with ‘large-scale experiments’; this will
give us important new insights and advance our
understanding of migration.
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References
Aarts, G., M., MacKenzie, B., McConnell, M., Fedak, and
Matthiopoulos, J. (2008) Estimating space-use and habitat preference from wildlife telemetry data. Ecography,
31, 140–60.
Able, K.P. (1982) Skylight polarization patterns at dusk
influence migratory orientation in birds. Nature, 299,
550–51.
Able, K.P. (1970) Radar study of altitude of nocturnal passerine migration. Bird-Banding, 41, 282–90.
Able, K.P. and Belthoff, J.R. (1998) Rapid ‘evolution’ of
migratory behavior in the introduced house finch of
eastern North America. Proceedings of the Royal Society
B, Biological Sciences, 265, 2063–71.
Abraham, E.R. (2007) Sea-urchin feeding fronts. Ecological
Complexity, 4, 161–68.
ACIA (2004) Impacts of a Warming Arctic—Arctic Climate
Impact Assessment, Cambridge University Press:
Cambridge.
Adams, C.C. (1918) Migration as a factor in evolution: its
ecological dynamics. The American Naturalist, 52,
465–90.
Adams, M.E. (1982) The Baggara problem: attempts at
modern change in Southern Darfur and Southern
Kordofan (Sudan). Development and Change, 13, 259–89.
Adriaensen, F., Ulenaers, P., and Dhondt, A.A. (1993) Ringing recoveries and the increase in numbers of European
great crested grebes (Podiceps cristatus). Ardea, 81, 59–70.
Ahlén, I., Hans, J. and Bach, L. (2009) Behavior of
Scandinavian bats during migration and foraging at
sea. Journal of Mammalogy, 90, 1318–23.
Ahola, M., Laaksonen, T., Sippola, K., Eeva, T., Rainio, K.,
and Lehikoinen, E. (2004) Variation in climate warming
along the migration route uncouples arrival and breeding dates. Global Change Biology, 10, 1610–17.
Åkesson, S. and Hedenström, A. (2007) How migrants get
there: migratory performance and orientation.
BioScience, 57, 123–33.
Åkesson, S. and Hedenström, A. (2000) Wind selectivity of
migratory flight departures in birds. Behavioral Ecology
and Sociobiology, 47, 140–44.
Åkesson, S., Broderick, A. C., Glen, F., Godley, B. J., Luschi,
P., Papi, F. and Hays, G. C. (2003) Navigation by green
216
turtles: which strategy do displaced adults use to find
Ascension Island? Oikos, 103, 363–72.
Albon, S.D. and Langvatn, R. (1992) Plant phenology and
the benefits of migration in a temperate ungulate. Oikos,
65, 502–13.
Alerstam, T. and Hedenström, A. (1998) The development
of bird migration theory. J. Avian. Biol., 29, 343–69.
Alerstam, T. and Pettersson, S.G. (1991) Orientation along
great circles by migrating birds using a sun compass.
Journal of Theoretical Biology, 152, 191–202.
Alerstam, T. (1978) Graphical illustration of pseudo-drift.
Oikos, 30, 409–12.
Alerstam, T. (1979) Wind as selective agent in bird migration. Ornis Scandinavica, 10, 76–93.
Alerstam, T. (1990) Bird migration, Cambridge University
Press: Cambridge.
Alerstam, T. (2001) Detours in bird migration. Journal of
Theoretical Biology, 209, 319–31.
Alerstam, T. (2006) Conflicting evidence about longdistance animal navigation. Science, 313, 791–94.
Alerstam, T. and Enckell, P.H. (1979) Unpredictable habitats
and evolution of bird migration. Oikos, 33, 228–32.
Alerstam, T. and Gudmundsson, G.A. (1999) Migration
patterns of tundra birds: tracking radar observations
along the Northeast Passage. Arctic, 52, 4, 346–71.
Alerstam, T. and Hedenström, A. (1998) The development
of bird migration theory. Journal of Avian Biology, 29,
343–69.
Alerstam, T. and Lindström, Å. (1990) Optimal bird migration: the relative importance of time, energy, and safety.
In E. Gwinner (ed.) Bird migration: Physiology and
Ecophysiology, pp. 331–51, Springer: Berlin.
Alerstam, T., Hedenström, A., and Åkesson, S. (2003)
Long-distance migration: evolution and determinants.
Oikos, 103, 247–60.
Alerstam, T., Gudmundsson, G.A., Green, M., and
Hedenström, A. (2001) Migration along orthodromic sun
compass routes by arctic birds. Science, 291, 300–303.
Alexander, R.M. (2000) Walking and running strategies for
humans and other mammals. In P. Domenici and R.W.
Blake (eds) Biomechanics in animal behaviour,
pp. 49–57, BIOS Scientific Publishers: Oxford.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Alexander, R.M. (2000) Principles of animal locomotion,
Princeton University Press: Princeton.
Alimaev, I.I. (2003) Transhumant ecosystems: fluctuations
in seasonal pasture productivity. In C. Kerven (ed.)
Prospects for Pastoralism in Kazakhstan and
Turkmenistan: From State Farms to Private Flocks,
pp. 31–51, RoutledgeCurzon: London.
Allen, C.R. and Saunders, D.A. (2002) Variability
between scales: predictors of nomadism in birds of an
Australian Mediterranean-climate ecosystem. Ecosystems, 5, 348–59.
Allen, C.R. and Saunders, D.A. (2006) Multimodel inference and the understanding of complexity, discontinuity, and nomadism. Ecosystems, 9, 694–99.
Ambrosini, R., MØller, A.P., and Saino, N. (2009) A quantitative measure of migratory connectivity. Journal of
Theoretical Biology, 257, 203–11.
Amstrup, S.C., McDonald, T.L, and Manly, B.F.J. (2005)
Handbook of Capture-Recapture Analysis, Princeton
University Press: Princeton, New Jersey.
Andersen, R. (1991) Habitat Deterioration and the
Migratory Behaviour of Moose (Alces alces L.) in Norway.
Journal of Applied Ecology, 28, 1, 102–108.
Anderson, J.H. and Quinn, T.P. (2007) Movements of adult
coho salmon (Oncorhynchus kisutch) during colonization
of newly accessible habitat. Canadian Journal of
Fisheries and Aquatic Sciences, 64, 8, 1143–54.
Anderson, W.B. and Polis, G.A. (1999) Nutrient fluxes
from water to land: seabirds affect plant nutrient status
on Gulf of California islands. Oecologia, 118, 324–32.
Andersson, M. (1980) Nomadism and site tenacity as alternative reproductive tactics in birds. Journal of Animal
Ecology, 49, 175–84.
Andreassen, H.P., Gundersen, H., and Storaas, T. (2005)
The effect of scent-marking, forest-clearing and supplemental feeding on moose-train collisions. Journal of
Wildlife Management, 69, 3, 1125–32.
Andriansen, H.K. (2003) The use and perception of mobility among Senegalese Fulani: New approaches to pastoral mobility, Kongevej Working Paper, Institute for
International Studies: Copenhagen.
Anstey, M.L., Rogers, S.M.,. Ott, S.R, Burrows, M., and
Simpson S.J. (2009) Serotonin mediates behavioral gregarization underlying swarm formation in desert
locusts. Science, 323, 627–30.
Arendt, J. and Reznick, D. (2008) Convergence and parallelism reconsidered: what have we learned about the genetics
of adaptation. Trends in Ecology and Evolution, 23, 26–32.
Arizaga, J., Barba, E., and Belda, E. J. (2008) Fuel management and stopover duration of blackcaps Sylvia atricapilla stopping over in northern Spain during autumn
migration period. Bird Study, 55, 124–34.
217
Armstrong, R.B. and Laughlin, M.H. (1985) Metabolic
indicators of fibre recruitment in mammalian muscles
during locomotion. Journal of Experimental Biology,
115, 201–13.
Arnason, R., Magnússon, G., and Agnarsson, S. (2000)
The Norwegian spring-spawning herring fishery: a stylized game model. Marine Resource Economics, 15,
293–319.
Apostolaki, P., Milner-Gulland, E.J., McAllister, M. and
Kirkwood, G. (2002) Modelling effects of establishing
marine reserves in nursery or spawning grounds.
Canadian Journal of Fisheries and Aquatic Science, 59,
405–15.
Aristotle (350 BC) Historia Animalium.
Arnold, S.J. (1983) Morphology, performance and fitness.
American Zoologist, 23, 347–61.
Auer, N.A. (1996) Importance of habitat and migration to
sturgeons with emphasis on lake sturgeon. Canadian
Journal of Fisheries and Aquatic Sciences, 53, Suppl. 1,
152–60.
Augustine, D.J. (2003) Long-term, livestock-mediated
redistribution of nitrogen and phosphorus in an East
African savanna. Journal of Applied Ecology, 40,
137–49.
Augustine, D.J., McNaughton, S.J., and Frank, D.A. (2003)
Feedbacks between soil nutrients and large herbivores
in a managed savanna ecosystem. Ecological Applications, 13, 1325–37.
Ayantunde, A.A. (1998) Influence of grazing regimes on
cattle nutrition and performance and vegetation dynamics in Sahelian rangelands, University of Wageningen:
Wageningen, The Netherlands.
Ayantunde, A.A., Fernández-Rivera, S., Hiernaux, P.H.Y.,
and van Keulen, H. (2001) Effect of timing and duration
of grazing of growing cattle in the West African Sahel
on diet selection, faecal output, eating time, forage
intake and live-weight changes. Animal Science, 72,
117–28.
Bäckman, J. and Alerstam, T. (2003) Orientation scatter of
free-flying nocturnal passerine migrants: components
and causes. Animal Behaviour, 65, 987–96.
Bailey, D.W., J.E. Gross, Laca, E.A., Rittenhouse, L.R.,
Coughenour, M.B., Swift, D.M., and Sims, P.L. (1996)
Mechanisms that result in large herbivore grazing distribution patterns. Journal of Range Management, 49, 5,
386–400.
Bairlein, F. (1998) The effect of diet composition on migratory fuelling in garden warblers Sylvia borin. Journal of
Avian Biology, 29, 546–51.
Bairlein, F. and Gwinner, E. (1994) Nutritional mechanisms
and temporal control of migratory energy accumulation
in birds. Annual Review of Nutrition, 14, 187–215.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
218
REFERENCES
Baker, A.J., González, P.M., Piersma, T., Niles, L.J., de
Lima Serrano do Nascimento, I., Atkinson, P.W., Clark,
N.A., Minton, C.D.T., Peck, M.K., and Aarts, G. (2004)
Rapid population decline in red knots: fitness consequences of decreased refueling rates and late arrival in
Deaware Bay. Proceedings of the Royal Society
B-Biological Science, 271, 875–82.
Baker, D.L. and Hobbs, N.T. (1985) Emergency feeding of
mule deer during winter: tests of a supplemental ration.
Journal of Wildlife Management, 49, 4, 934–42.
Baker, R.R. (1978) The Evolutionary Ecology of Animal
Migration, Hodder and Stoughton: London.
Balmford, A., Leader-Williams, N. and Green, M.J.B. (1995)
Parks or arks: where to conserve threatened mammals?
Biodiversity and Conservation, 4, 595–607.
Banko, W. (1960) The Trumpeter Swan, North American
Fauna, No. 63, U.S. Fish and Wildlife Service:
Washington, DC.
Barnes, R.F.W. (1999) Is there a future for elephants in
West Africa? Mammal Review, 29, 3, 175–99.
Barnett-Johnson, R., Ramos, F.C., Grimes, C.B., and
MacFarlane, R.B. (2005) Validation of Sr isotopes in otoliths by laser ablation multicollector inductively coupled
plasma mass spectrometry (LA-MC-ICPMS): opening
avenues in fisheries science applications. Canadian
Journal of Fisheries and Aquatic Sciences, 62, 2425–30.
Barta. Z, McNamara, J.M., Houston, A.I., Weber, T.P.,
Hedenström, A., and Féro, O. (2008) Optimal moult
strategies in migratory birds. Philosophical Transactions
of the Royal Society of London B, 363, 211–29.
Bartumeus, F. and Levin, S.A. (2008) Fractal reorientation
clocks: Linking animal behavior to statistical patterns of
search. Proceeding of the National Academy of Sciences
USA, 105, 19072–77.
Bartumeus, F., Da Luz, M.G.E., Viswanathan, G.M., and
Catalan, J. (2005) Animal search strategies: a quantitative random-walk analysis. Ecology, 86, 3078–87.
Baskin, L. (1991) Reindeer husbandry in the Soviet Union.
In L. A. Renecker and R. J. Hudson (eds) Wildlife
Production: Conservation and Sustainable Development,
pp. 218–26, Agricultural and Forestry Experiment
Station: Fairbanks, Alaska.
Baskin, Y. (1993) Trumpeter swans relearn migration.
BioScience, 43, 2, 76–9.
Bassett, T.J. (1986) Fulani herd movements. The
Geographical Review, 76, 233–48.
Bassett, T.M. and Turner, M.D. (2007) Sudden shift or
migratory drift? Fulbe herd movements to the SudanoGuinean region of West Africa. Human Ecology, 35,
33–49.
Batima, P. (2006) Climate change vulnerability and adaptation in the livestock sector of Mongolia: A final report
submitted to Assessments of Impacts and Adaptations
to Climate Change (AIACC), Project No. AS 06.
International START Secretariat: Washington, D.C.
Bauchinger, U. and Klaassen, M. (2005) Longer days in
spring than in autumn accelerate migration speed.
J. Avian Biol., 36, 3–5.
Bauchinger, U. and McWilliams, S. (2009) Carbon turnover in tissues of a passerine bird: Allometry, isotopic
clocks, and phenotypic flexibility in organ size.
Physiological and Biochemical Zoology, 82, 787–97.
Bauer, S., Gienapp, P., and Madsen, J. (2008) The relevance of
environmental conditions for departure decision changes
en route in migrating geese. Ecology, 89, 1953–60.
Bauer, S., Barta, Z., Ens, B.J., Hays, G.C., McNamara, J.M.,
and Klaassen, M. (2009) Animal migration: linking
models and data beyond taxonomic limits. Biology
Letters, 5, 433–35.
Bayer, W. and Waters-Bayer, A. (1994) Forage alternatives
from range and field: pastoral forage management and
improvement in the African drylands. In I. Scoones (ed.)
Living with Uncertainty: New Directions in Pastoral
Development in Africa, pp. 58–78, Intermediate
Technology Publications Ltd: London.
Bayly, N.J. (2006) Optimality in avian migratory fuelling
behaviour: a study of a trans-Saharan migrant. Animal
Behaviour, 71, 173–82.
Bayly, N.J. (2007) Extreme fattening by sedge warblers,
Acrocephalus schoenobaenus, is not triggered by food
availability alone. Animal Behaviour, 74, 471–79.
Bazazi, S., Buhl, J., Hale, J.J., Anstey, M.L., Sword, G.A.,
Simpson, S.J., and Couzin, I.D. (2008) Collective motion
and cannibalism in locust migratory bands. Current
Biology, 18, 735–39.
Bazazi S., Romanczuk, P., Thomas, S., Schimansky-Geier,
L., Hale, J.J., Miller, G.A., Sword, G.A., Simpson, S.J.,
and Couzin, I.D. (in press) Nutritional state and
collective motion: from individuals to mass migration.
Proceedings of the Royal Society B, Biological Sciences.
Beach, H. (1981) Reindeer-herd management in transition:
the case of Tuorpon Saameby in Northern Sweden,
Uppsala Studies in Cultural Anthropology 3:
Stockholm.
Beach H. and Stammler, F. (2006) Human–Animal relations in pastoralism. In F. Stammler and H. Beach (eds)
People and Reindeer on the Move, Special Issue of the
journal Nomadic Peoples,10, 2, pp. 5–29, Berghahn
Publishers: Oxford.
Bearhop, S., Fiedler, W., Furness, R.W., Votier, S.C.,
Waldron, S., Newton, J., Bowen, G.J., Berthold, P., and
Farnsworth, K. (2005) Assortative mating as a mechanism for rapid evolution of a migratory divide. Science,
310, 502–504.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Beauvilain, A. (1977) Les Peul du Dallol Bosso, Études
Nigériennes, 42, Institut de recherche en sciences
humaines: Niamey, Niger.
Bedard, J., Nadeau, A., and Gauthier, G. (1986) Effects of
spring grazing by greater snow geese on hay production. Journal of Applied Ecology, 23, 65–75.
Bedunah, D. and Harris, R. (2005) Observations on changes
in Kazak pastoral use in two townships in Western
China: A loss of traditions. Nomadic Peoples, 9, 107–29.
Beekman J.H., Nolet B.A., and Klaassen, M. (2002)
Skipping swans: Fuelling rates and wind conditions
determine differential use of migratory stopover sites of
Bewick’s Swans Cygnus bewickii. Ardea, 90, 437–59.
Beekman, J.H., van Eerden, M.R. and Dirksen, S. (1991)
Bewick’s swans Cygnus columbianus bewickii utilising
the changing resource of Potamogeton pectinatus during
autumn in the Netherlands. Wildfowl, Suppl. 1,
238–48.
Behnke, R.H. (1994) Natural resource management in pastoral Africa. Development Policy Review, 12, 1, 5–27.
Behnke, R.H. (1999) Stock movement and range-management in a Himba community in north-western Namibia.
In M. Niamir-Fuller (ed.) Managing Mobility in African
Rangelands: the Legitimization of Transhumance,
pp. 184–216, Intermediate Technology Publications:
London.
Behnke, R.H. (2008) The drivers of fragmentation in arid
and semi-arid landscapes. In Galvin, K.A., Reid, R.S.,
Behnke, R.H., and Hobbs, N.T. (eds) Fragmentation in
Semi-Arid and Arid Landscapes: Consequences for
Human and Natural Systems, pp. 305–40, Dordrecht,
The Netherlands: Springer.
Behnke, R.H. and Scoones, I. (1993) Rethinking range
ecology: implications for rangeland management in
Africa. In R.H. Behnke, I. Scoones, and C. Kerven (1993)
Range Ecology at Disequilibrium: New Models of
Natural Variability and Pastoral Adaptation in African
Savannas, pp. 1–30, Overseas Development Institute:
London.
Behnke, R.H., Davidson, G., Jabbar, A., and Coughenour,
M. (2008) Human and natural factors that influence livestock distributions and rangeland desertification in
Turkmenistan. The Socio-Economic Causes and
Consequences of Desertification and Central Asia, pp.
141–68, Springer Science + Business Media B.V.:
Dordrecht, The Netherlands.
Bekenov, A.B., Grachev, Iu. A., and Milner-Gulland, E.J.
(1998) The ecology and management of the saiga antelope in Kazakhstan. Mammal Review, 28, 1–52.
Bell, C.P. (2000) Process in the evolution of bird migration
and pattern in avian ecogeography. Journal of Avian
Biology, 31, 258–65.
219
Bell, C.D., Blumenthal, J.M., Austin, T.J., Solomon J.L.,
Ebanks-Petrie, G., Broderick, A.C., and Godley, B.J.
(2006) Traditional Caymanian fishery may impede local
marine turtle population recovery. Endangered Species
Research, 2, 63–69.
Bell, C., Solomon, J.L., Blumenthal, J.M., Austin, T.J., EbanksPetrie, G., Broderick, A.C., and Godley, B.J. (2007)
Monitoring and conservation of critically reduced marine
turtle nesting populations: lessons from the Cayman
Islands. Animal Conservation, 10, 39–47.
Bellrose, F.C. (1958) The orientation of displaced waterfowl in migration. Wilson Bull., 70, 20–40.
Bengis, R.G., Grant, R., and de Vos, V. (2003) Wildlife diseases and veterinary controls: a savannah ecosystem
perspective. In J. Du Toit, K.H. Rogers, and H.C. Biggs
(eds) The Kruger Experience, pp. 349–69, Island Press:
Washington.
Benhamou, S. (2006) Detecting an orientation component
in animal paths when the preferred direction is individual-dependent. Ecology, 87, 518–28.
Benoit, M. (1979) Le chemin des Peuls du Boobola: contribution à l’écologie du pastoralisme en Afrique des
savanes. O.R.S.T.O.M.: Paris.
Berg, H. (1983) Random walks in biology, Princeton
University Press: Princeton.
Berger, J. (1998) Future prey: some consequences of the
loss and restoration of large carnivores. In T.M. Caro
(ed.) Behavioral Ecology and Conservation Biology,
pp. 80–100, Oxford University Press: New York.
Berger, J. (2004) The last mile: how to sustain long-distance
migration in mammals. Conservation Biology, 18, 2,
320–31.
Berger, J., Cain, S.L., and Berger, K.M. (2006) Connecting
the dots: an invariant migration corridor links the
Holocene to the present. Biol. Letters, 2, 528–31.
Berkes, F. (1999) Sacred Ecology: Traditional Ecological
Knowledge and Resource Management, Taylor and
Francis: Philadelphia and London.
Berlioz, J. (1950) Carreteres generaux et origines des
migrations. In P Grasse (ed.) Trate de zoologie, Vol. XV,
Oiseaux, pp. 1074–88.
Bernhardt, E.S., Palmer, M.A., Allan, J.D., Alexander,
G., Barnas, K., Brooks, S., Carr, J., Clayton, S., Dahm,
C., Follstad-Shah, J., Galat, D., Gloss, S., Goodwin, P., Hart,
D., Hassett, B., Jenkinson, R., Katz S., Kondolf, G.M., Lake,
P.S., Lave, R., Meyer, J.L., O’Donnell, T.K., Pagano, L.,
Powell, B., and Sudduth, E. (2005) Synthesizing U.S. river
restoration efforts. Science, 308, 636–37.
Berthold, P. (1984) The control of partial migration in birds:
a review. Ring, 10, 253–65.
Berthold, P. (1991) Genetic control of migratory behaviour
in birds. Trends in Ecology and Evolution, 6, 8, 254–57.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
220
REFERENCES
Berthold, P. (1996) Control of Bird Migration, Chapman
and Hall: London.
Berthold, P. (1999) A comprehensive theory for the evolution, control and adaptability of avian migration.
Ostrich, 70, 1–11.
Berthold, P. (2001) Bird Migration. A General Survey,
Oxford University Press: Oxford.
Berthold, P. and Querner, U. (1981) Genetic basis of migratory behavior in European warblers. Science, 212,
77–79.
Berthold, P., van den Bossche, W., Fiedler, W., Gorney,
E., Kaatz, M., Leshem, Y., Nowak, E., et al. (2001) The
migration of the White Stork (Ciconia ciconia): a special
case according to new data. Journal für Ornithologie,
142, 73–92.
Bevan, R.M., Butler, P.J., Woakes, A.J., and Prince, P.A.
(1995) The energy expenditure of free-ranging blackbrowed albatrosses. Philosophical Transactions of the
Royal Society B, 350, 119–31.
Biebach, H. (1983) Genetic determination of partial migration in the European robin (Erithacus rubecula). Auk, 100,
601–606.
Biebach, H. (1998) Phenotypic organ flexibility in Garden
Warblers Sylvia borin during long-distance migration.
Journal of Avian Biology, 29, 529–35.
Biebach, H., Friedrich, W., and Heine, G. (1986) Interaction
of body mass, fat, foraging and stopover period in transSahara migrating passerine birds. Oecologia, 69, 370–79.
Birdlife International (2004) Tracking ocean wanderers:
the global distribution of albatross and petrels,
Results from the Global Procellariiform tracking
Workshop, 1–5 September, 2003, Birdlife International:
Cambridge, UK.
Bishop, C.M. and Butler, P.J. (1995) Physiological modeling
of oxygen-consumption in birds during flight. Journal
of Experimental Biology, 198, 2153–63.
Bishop, J., McKay, H., Parrott, D., and Allan, J. (2003)
Review of international research literature regarding
the effectiveness of auditory bird scaring techniques
and potential alternatives. UK Department for
Environment, Food and Rural Affairs Report: London.
Bishop, C.M., Ward, S., Woakes, A.J., and Butler, P.J. (2002)
The energetics of barnacle geese (Branta leucopsis) flying
in captive and wild conditions. Comparative
Biochemistry and Physiology a-Molecular and
Integrative Physiology, 133, 225–37.
Bisson, I.A., Safi, K., and Holland, R.A. (2009) Evidence for
repeated independent evolution of migration in the
largest family of bats. PLoS ONE, 4, e7504.
Biuw, E., Boehme, L., Guinet, C., Hindell, M., Costa,
D., Charrassin, J-B., Roquet, F., et al. (2007) Variations in
behavior and condition of a Southern Ocean top preda-
tor in relation to in situ oceanographic conditions.
Proceedings of the National Academy of Sciences of the
United States of America, 104, 13705–10.
Bjorndal, K.A. (1997) Foraging ecology and nutrition of
sea turtles. In P.L. Lutz and J.A. Musick (eds) The
Biology of Sea Turtles, pp. 199–231, CRC Press:
London.
Blackwell, P.G. (1997) Random diffusion models for animal movement. Ecological Modelling, 100, 87–102.
Blackwell, P.G. (2003) Bayesian inference for Markov processes with diffusion and discrete components.
Biometrika, 90, 613–27.
Blaxter, K.L., Wainman, F.W., and Wilson, R. (1961) The
regulation of food intake in sheep. Journal of Animal
Production, 3, 51–61.
Bleich, V.C. and Pierce, B.M. (2001) Accidental mass mortality of migrating mule deer. Western North American
Naturalist, 61, 1, 124–25.
Blem, C.R. (1980) The energetics of migration. In
S.A. Gauthreaux (ed.) Animal Migration, Orientation
and Navigation, pp. 175–224, Academic Press: London.
Block, B.A., Teo, S.L.H. Walli, A. Boustany, A. Stokesbury,
M.J.W. Farwell, C.J. Weng, K.C., Dewar, H., and
Williams, T.D. (2005) Electronic tagging and population
structure of Atlantic bluefin tuna. Nature, 434, 1121–27.
Block, W.M., Franklin, A.B., Ward, J.P. Jr., Ganey, J.I.,
and White, G.C. (2001) Design and implementation
of monitoring studies to evaluate the success of ecological restoration on wildlife. Restoration Ecology, 9,
3, 293–303.
Blumstein, D.T. (2002) Moving to suburbia: ontogenetic
and evolutionary consequences of life on predator-free
islands. Journal of Biogeography, 29, 685–92.
Bolger, D.T., Newmark, W.D., Morrison, T.A., and Doak,
D.F. (2008) The need for integrative approaches to
understand and conserve migratory ungulates. Ecology
Letters, 11, 63–77.
Bolker, B.M. 2008, Ecological Models and Data in
R, Princeton University Press: Princeton.
Bollens, S.M. and Frost, B.W. (1989) Predator-induced diel
vertical migration in a planktonic copepod. Journal of
Plankton Research, 11, 1047–65.
Bollens, S.M. and Frost, B.W. (1989) Zooplanktivorous
fish and variable diel migration in the marine plankt
onic copepod Calanus pacificus. Limnology and
Oceanography, 34, 1072–83.
Bonfiglioli, Angelo Maliki (1988) Dudal: Histoire de famille
et histoire de troupeau chez un groupe de Wodaabe du
Niger, Cambridge University Press: Cambridge.
Boone, R.B., Thirgood, S.J., and Hopcraft, J.G.C. (2006)
Serengeti wildebeest migratory patterns modeled from
rainfall and new vegetation growth. Ecology, 87, 1987–94.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Börger, L., Dalziel, B.D., and Fryxell, J.M. (2008) Are there
general mechanisms of animal home range behaviour?
A review and prospects for future research. Ecology
Letters, 11, 637–50.
Both, C. and Visser, M.E. (2001) Adjustment to climate
change is constrained by arrival date in a long-distance
migrant bird. Nature, 411, 296–98.
Both, C., Bouwhuis, S., Lessells, C.M., and Visser, M.E.
(2006) Climate change and population declines in a
long-distance migratory bird. Nature, 441, 81–83.
Bowlin, M.S. and Wikelski, M.C. (2006) Calibration of
heart rate and energy expenditure during flight and at
rest in a passerine. Integrative and Comparative Biology,
46, E14–E14.
Bowlin, M. S. and Wikelski, M. (2006) Pointed wings, low
wingloading and calm air reduce migratory flight costs
in songbirds. PLoS ONE, 3, e2154.
Bowlin, M. S., Cochran, W.W., and Wikelski, M.C. (2005)
Biotelemetry of New World thrushes during migration:
Physiology, energetics and orientation in the wild.
Integrative and Comparative Biology, 45, 295–304.
Boyd, I.L. (2004) Migration of marine mammals. In
D. Werner (ed.) Biological Resources and Migration, pp
203–10, Springer-Verlag: Berlin.
Boyle, P, and Rodhouse, P. (2005) Cephalopods: ecology
and fisheries, Blackwell: London.
Boyle, W.A. and Conway, C.J. (2007) Why migrate? A test
of the evolutionary precursor hypothesis. The American
Naturalist, 169, 344–59.
Bradshaw, G.A. and Spies, T.A. (1992) Characterizing canopy gap structure in forests using wavelet analysis.
Journal of Ecology, 80, 205–15.
Bradshaw, W.E. and Holzapfel, C.M. (2006) Evolutionary
response to rapid climate change. Science, 312, 1477–78.
Brattström, O., Wassenaar, L. I., Hobson, K. A., and
.
Akesson, S. (2008) Placing butterflies on the map—testing regional geographical resolution of three stable isotopes in Sweden using the monophagus peacock Inachis
io. Ecography, 31, 490–98.
Breiman, L. (2001) Statistical modeling: the two cultures.
Statistical Science, 16, 199–231.
Breman, H. and de Wit, C.T. (1983) Rangeland productivity and exploitation in the Sahel. Science, 221, 1341–47.
Broderick A.C., Coyne, M.C., Fuller, W.J., Glen, F., and
Godley, B.J. (2007) Fidelity and over-wintering of sea
turtles. Proceedings of the Royal Society of London
Series B, 274, 1533–38.
Broderick, A.C., Frauenstein, R., Glen, F., Hays, G.C.,
Jackson, A.L., Pelembe, T., Ruxton, G.D., and Godley,
B.J. (2006) Are green turtles globally endangered? Global
Ecology and Biogeography, 15, 21–6. www.seaturtle.
org/mtrg/pubs/Broderick_GEB_2006.pdf.
221
Brönmark, G., Skov, C., Broderson, J., Nilsson, P.A., and
Hansson, L.-A. (2008) Seasonal migration determined
by a trade-off between predator avoidance and growth.
PLoS ONE, 3, e1957.
Brooks, T.M., Mittermeier, R.A., Mittermeier, C.G. da
Fonseca, G.A.B., Rylands, A.B., Konstant, W.R., Flick,
P., Pilgrim, J., Oldfield, S., Magin, G., and Hilton-Taylor,
C. (2002) Habitat loss and extinction in the hotspots of
biodiversity. Conservation Biology, 16, 4, 909–23.
Brower, L.P. and Malcolm, S.B. (1991) Animal Migrations:
Endangered Phenomena. American Zoologist, 31, 1,
265–76.
Brower, L. P., Fink, L. S., and P. Walford (2006) Fueling the
fall migration of the monarch butterfly. Integrative and
Comparative Biology, 46, 1123–42.
Brown, C.G. (1992) Movement and migration patterns of
mule deer in southeastern Idaho. Journal of Wildlife
Management, 56, 246–53.
Brown, C.G., Waldron, S.A., and Longworth, J.W. (2008)
Sustainable Development in Western China: Managing
People, Livestock and Grasslands in Pastoral Areas,
Edward Elgar: Cheltenham, UK.
Bruderer, B. and Salewski, V. (2008) Evolution of bird
migration in a biogeographical context. Journal of
Biogeography, 35, 1951–59.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L.,
Borchers, D.L., and Thomas, L. (2001) Introduction to
Distance Sampling: Estimating Abundance of Biological
Populations, Oxford University Press: Oxford.
Buckland, S.T., Anderson, D. R., Burnham, K. P., Laake, J.
L., Borchers, D. L. and Thomas L. (2004) Advanced
Distance Sampling: Estimating Abundance of Biological
Populations, Oxford University Press: Oxford.
Buhl, J., Sumpter, D.J.T., Couzin, I.D., Hale, J.J., Despland,
E., Miller, E.R. and Simpson, S.J. (2006) From disorder to
order in marching locusts. Science, 312, 1402–406.
Burnham, K.P. and D.R. Anderson. 2002, Model
Selection and Multi-Model Inference, A Practical
Information–Theoretic Approach, Springer Verlag:
Berlin, Germany.
Butler, P.J. (1982) Respiration during flight and diving in
birds. In A.D.F. Addink and N. Spronk (eds) Exogenous
and Endogenous Influences on Metabolic and Neural
Control, pp 103–14, Pergamon Press: London.
Butler, P.J. (1986) Exercise. In S. Nilsson and S. Holmgren
(eds) Fish Physiology: Recent Advances, pp. 102–18,
Croom Helm: London.
Butler, P.J. (1991) Exercise in birds. Journal of Experimental
Biology, 160, 233–62.
Butler, P.J. and Bishop, C.M. (2000) Flight. In G. C. Whittow
(ed.) Sturkie’s Avian Physiology, 5th edn, Academic
Press: New York, NY.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
222
REFERENCES
Butler, P.J. and Woakes, A.J. (1980) Heart-rate, respiratory
frequency and wing beat frequency of free flying barnacle geese Branta leucopsis. Journal of Experimental
Biology, 85, 213–26.
Butler, P.J., Woakes, A.J., and Bishop, C.M. (1998) Behaviour
and physiology of Svalbard barnacle geese Branta leucopsis during their autumn migration. Journal of Avian
Biology, 29, 536–45.
Bykova, E., Esipov, A., and Chernogaev, E. (2009) Will the
saiga return to its traditional breeding grounds in
Uzbekistan? Saiga News, 8, 13–5. [Online], Available:
http://www.saiga-conservation.com/saiga_news.
html. [April 2009].
Carmi, N., Pinshow, B., Porter, W.P., and Jaeger, J. (1992)
Water and energy limitations on flight duration in small
migrating birds. Auk, 109, 268–76.
Caro, T.M., Pelkey, N., Borner, M., Campbell, K.L.I.,
Woodworth, B.L., Farm, B.P., Kuwai, J.O., Huish, S.A., and
Severre, E.L.M. (1998) Consequences of different forms of
conservation for large mammals in Tanzania: preliminary
analyses. African Journal of Ecology, 36, 303–20.
Carpenter, F.L., Hixon, M.A., Russell, R.W., Paton, D.C.,
and Temeles E. J. (1993) Interference asymmetries
among age-sex classes of rufous hummingbirds during
migratory stopovers. Behavioral Ecology and
Sociobiology, 33, 297–304.
Carr, A. (1984) The sea turtle: so excellent a fishe. University
of Texas Press: Austin.
Chaloupka, M., Bjorndal, K.A., Blalzs, G.H., Bolten, A.B.,
Ehrhart, L.M., Limpus, C.J., Suganuma, H., Troeng, S.,
and Yamaguchi, M. (2008) Encouraging outlook for
recovery of a once severely exploited marine megaherbivore. Global Ecology and Biogeography, 17, 2,
297–304.
Chambers, L.E. (2008) Trends in timing of migration of
south-western Australian birds and their relationship to
climate. Emu, 108, 1–14.
Chan, K. (2001) Partial migration in Australian landbirds:
a review. Emu, 101, 281–92.
Chapin III, F.S., Zavaleta, E.S. Eviner, V.T. Naylor, R. L.
Vitousek, P.M. Reynolds, H.L. Hooper, D.U. Lavorel, S.
Sala, O.E. Hobbie, S.E. Mack M.C., and Díaz, S. (2000)
Consequences of changing biodiversity. Nature, 405,
234–42.
Chapman, J.W., Reynolds, D.R., Brooks, S.J., Smith, A.D.,
and Woiwod, I.P. (2006) Seasonal variation in the migration strategies of the green lacewing Chrysoperla carnea
species complex. Ecological Entomology, 31, 378–88.
Chapman, J.W., Reynolds, D.R., Smith, A.D., Smith, E.T.,
and Woiwod, I.P. (2004) An aerial netting study of
insects migrating at high altitude over England. Bulletin
of Entomological Research, 94, 123–36.
Chapman, J.W., Reynolds, D.R., Hill, J.K., Sivell, D., Smith,
A.D., and Woiwod, I.P. (2008a) A seasonal switch in
compass orientation in a high-flying migrant moth.
Current Biology, 18, R908–R909.
Chapman, J.W., Reynolds, D.R., Mouritsen, H., Hill,
J.K., Riley, J.R., Sivell, D., Smith, A. D., and Woiwod, I.P.
(2008b) Wind selection and drift compensation optimize
migratory pathways in a high-flying moth. Current
Biology, 18, 514–18.
Cheke, R.A. and Tratalos, J.A. (2007) Migration, patchiness, and population processes illustrated by two
migrant pests. Bioscience, 57, 145–54.
Cherel, Y., Kernaléguen, L., Richard, P., and Guinet,
C. (2009) Whisker isotopic signature depicts migration
patterns and multi-year intra- and inter-individual foraging strategies in fur seals. Biology Letters, 5, 830–32.
Chesser, R.T. and Levey, D.J. (1998) Austral Migrants and
the Evolution of Migration in New World Birds: Diet,
Habitat, and Migration Revisited. The American
Naturalist, 152, 311–19.
Chimeddorj, B. (2009) Public Awareness Needs Assessment
for the Saiga Antelope Conservation Project in Mongolia.
Saiga News, 8, 9–10. [Online], http://www.saigaconservation.com/saiga_news.html [April 2009].
Christie, K.S. and Reimchen, T.E. (2005) Post-reproductive
Pacific salmon, Oncorhynchus spp., as a major nutrient
source for large aggregations of gulls, Larus spp.
Canadian Field-Naturalist, 119, 202–207.
Ciancio, J.E., Pascual, M., Botto. M., Amaya, M., O’Neal, S.,
Riva Rossi, C., and Iribarne, O. (2008) Stable isotope profiles of partially migratory salmonid populations in Atlantic
rivers of Patagonia. Journal of Fish Biology, 72, 1708–19.
Cimprich, D.A. and F.R. Moore (2006) Fat affects predatoravoidance behavior in gray catbirds (Dumetella carolinensis) during migratory stopover. Auk, 123, 1069–76.
Cimprich, D.A., Woodrey, M.S., and Moore, F.R. (2005)
Passerine migrants respond to variation in predation
risk during stopover. Animal Behaviour, 69, 1173–79.
Cissé, S. (1981) L’avenir du pastoralisme dans le delta central du Niger (Mali): Agriculture, élevage, ou agropastoralisme? Nomadic Peoples, 9, 16–21.
CITES (Convention on International Trade in Endangered
Species of Wild Fauna and Flora) (2004) Conservation of
Saiga tatarica. CoP 13, Document 32. Convention on
International Trade in Endangered Species of Flora and
Fauna. [Online], http://www.cites.org/eng/cop/13/
doc/E13–32.pdf [April 2009].
Clark, C.W. and Mangel, M. (1986)The evolutionary advantages of group foraging. Theor. Pop. Biol., 30, 45–75.
Clark, C.W. and Mangel, M. (2000) Dynamic state variable
models in ecology: methods, and applications, Oxford
University Press: Oxford.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Clausen, P., Green, M., and Alerstam, T. (2003) Energy
limitations for spring migration and breeding: the case
of Brent geese Branta bernicla tracked by satellite telemetry to Svalbard and Greenland. Oikos, 103, 426–45.
Cleaveland, S., Packer, C., Hampson, K., Kaare, M., Kock,
R., Craft, M., Lembo, T., Mlengeya, T., and Dobson, A.
(2008) The multiple roles of infectious disease in the
Serengeti ecosystem. In A.R.E. Sinclair, C. Packer, S.A.R.
Mduma, and J.M. Fryxell (eds) Serengeti III: Human
Impacts on Ecosystem Dynamics, pp. 209–40 Chicago
University Press: Chicago.
Clevenger, A.P. and Waltho, N. (2005) Performance indices
to identify attributes of highway crossing structures
facilitating movement of large mammals. Biological
Conservation, 121, 453–64.
Clewett, J.F., Clarkson, N.M., George, D.A., Ooi, S.H.,
Owens, D.T., Partridge, I.J., and Simpson, G.B. (2003)
Rainman StreamFlow version 4.3: a comprehensive climate and streamflow analysis package on CD to assess
seasonal forecasts and manage climate risk, QI03040,
Department of Primary Industries: Queensland.
Clobert, J., Le Galliard, J. F., Cote, J., Meylan, S., and Massot
M. (2009) Informed dispersal, heterogeneity in animal
dispersal syndromes and the dynamics of spatially
structured populations. Ecology Letters, 12, 197–209.
Clutton-Brock, T., Guiness, F.E., and Albon, S.D. (1982)
Red Deer: Behavior and Ecology of Two Sexes, Wildlife
Behavior and Ecology Series, The University of Chicago
Press: Chicago.
CMS (Convention on Migratory Species) (2006a) Revised
Overview Report of the First Meeting of the Signatories
to the Memorandum of Understanding concerning
Conservation, Restoration and Sustainable Use of the
Saiga Antelope (Saiga tatarica tatarica) (CMS/SA- 1/
Report Annex 5), Convention on Migratory Species.
[Online], Available: http://www.cms.int/species/
saiga/post_session/Annex_05_Revised_Overview_
Report_E.pdf
CMS (Convention on Migratory Species) (2006b) Medium
term international work programme for the saiga antelope (2007–2011). (CMS/SA- 1/Report Annex 9):
Convention on Migratory Species. [Online], Available:
http://www.cms.int/species/saiga/post_session/
Annex_09_MediumTerm_Int_WrkProgm_E.pdf
Cochran, W.W. (1972) Long-distance tracking of birds. In
S.R Galler, K. Schmidt-Koenig, G.J. Jacobs, and R.E
Belleville (eds) Animal Orientation and Navigation, pp.
39–59, National Aeronautics and Space Adminstration:
Washington, DC.
Cochran, W.W. and Kjos, C.G. (1985) Wind drift and
migration of thrushes: A telemetry study. Illinois Natural
History Survey Bulletin, 33, 297–330.
223
Cochran, W.W. and Wikelski, M. (2005) Individual migratory tactics of New World Catharus thrushes: Current
knowledge and future tracking options from space. In
R. Greenberg and P.P. Marra (eds) Birds of Two Worlds:
The Ecology and Evolution of Migration, pp. 274–89,
John Hopkins University Press: Baltimore.
Cochran, W.W., Mouritsen, H., and Wikelski, M. (2004)
Migrating songbirds recalibrate their magnetic compass
daily from twilight cues. Science, 304, 405–408.
Codling, E.A., Pitchford, J.W., and Simpson, S.D. (2007)
Group navigation and the ‘many-wrongs principle’ in
models of animal movement. Ecology, 88, 1864–70.
Cohen, D. (1967) Optimization of seasonal migratory
behaviour. The American Naturalist, 101, 5–17.
Colin de Verdière, P. (1995) Étude compare de trois systèmes agropastoraux dans la région de Filingué—Niger.
Doctorate thesis, Université de Hohenheim Allemagne
et Institut National Agronomique: Paris-Grignon,
France.
Conover, M.R. (1984) Comparative effectiveness of Avitrol,
exploders and hawk-kites in reducing blackbird damage
to corn. Journal of Wildlife Management, 48, 1, 109–16.
Conradt, L., Clutton-Brock, T. H., and Thomson, D. (1999)
Habitat segregation in ungulates: are males forced into
suboptimal foraging habitats through indirect competition by females? Oecologia, 119, 367–77.
Conradt, L., Krause, J., Couzin, I. D., and Roper, T. J. (2009)
‘Leading according to need’ in self-organizing groups.
American Naturalist, 173, 304–12.
Cooke, S.J., Hinch, S.G., Wikelski, M., Andrews, R.D.,
Kuchel, L.J., Wolcott, T.G., and Butler P.J. (2004)
Biotelemetry: a mechanistic approach to ecology. Trends
in Ecology and Evolution, 19, 334–43.
Coppock, D.L., Ellis, J.E., and Swift, D.M. (1986) Livestock
feeding ecology and resource utilization in a nomadic
pastoral ecosystem. Journal of Applied Ecology, 23,
573–83.
Coppolillo, P.B. (2000) The landscape ecology of pastoral
herding: spatial analysis of land use and livestock
production in East Africa. Human Ecology, 28, 4,
527–60.
Coppolillo, P.B. (2001) Central-place analysis and modeling
of landscape-scale resource use in an East African agropastoral system. Landscape Ecology, 16, 205–19.
Coughenour, M.B. (1991) Spatial components of plantherbivore interactions in pastoral, ranching and native
ungulate ecosystems. Journal of Range Management,
44, 530–42.
Coughenour, M.B., Coppock, D.L., and Ellis, J.E. (1990)
Herbaceous forage variability in an arid pastoral region
of Kenya: importance of topographic and rainfall gradients. Journal of Arid Environments, 19, 147–59.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
224
REFERENCES
Coulson, T., Guinness, F., Pemberton, J., and Clutton-Brock
T. (2004) The demographic consequences of releasing a
population of red deer from culling. Ecology, 85, 411–22.
Courchamp, F., Berec, L., and Gascoigne, J. (2008) Allee
effects in ecology and conservation. Oxford University
Press: Oxford.
Coutant, C.C. and Whitney, R.R. (1999) Fish behavior in
relation to passage through hydropower turbines: a
review. Transactions of the American Fisheries Society,
129, 351–80.
Couzin, I.D. and Krause, J. (2003) Self-organization and
collective behavior in vertebrates. Advances in the
Study of Behavior, 32, 32, 1–75.
Couzin, I.D. and Laidre, M.E. (2009) Fission–fusion populations. Current Biology, 19, R633–35.
Couzin, I.D., Krause, J., Franks, N.R., and Levin S.A. (2005)
Effective leadership and decision-making in animal
groups on the move. Nature, 433, 513–16.
Cox, D.R. (2006) Principles of Statistical Inference,
Cambridge University Press: Cambridge, UK.
Cox, G.W. (1968) The role of competition in the evolution
of migration. Evolution, 22, 180–92.
Cox, G.W. (1985) The evolution of avian migration systems between temperature and tropical regions of the
New World. American Naturalist, 126, 451–74.
Cox, T.M., Lewison, R.I., Zydelis, R., Crowder L.B., Safina,
C., and Read, A.J. (2007) Comparing effectiveness of
experimental and implemented bycatch reduction
measures: the ideal and the real. Conservation Biology,
21, 1155–64.
Craig, A.S. and Herman, L.I. (1997) Sex differences in site
fidelity and migration of humpback whales (Megaptera
novaeangliae) to the Hawaiian Islands. Canadian Journal
of Zoology, 75, 1923–33.
Crick, H.Q.P. (2004) The impact of climate change on
birds. Ibis, 146, 48–56.
Cumming, H.G. and Beange, D.B. (1987) Dispersion and
movements of Woodland Caribou near Lake Nipigon,
Ontario. Journal of Wildlife Management, 51, 1, 69–79.
Cunnison, I. (1966) Baggara Arabs: Power and the Lineage
in a Sudanese Nomad Tribe, Clarendon Press: Oxford.
Curio, E., Ernst, U., and Vieth, W. (1978) Cultural transmission of enemy recognition: one function of mobbing.
Science, 202, 899–901.
Dalziel, B.D., Morales, J.M., and Fryxell, J.M. (2008) Fitting
probability distributions to animal movement trajectories:
using artificial neural networks to link distance, resources,
and memory. American Naturalist, 172, 248–58.
Darby, W.R. and Pruitt, W.O. (1984) Habitat use, movements and grouping behavior of Woodland Caribou,
Rangifer tarandus caribou, in southeastern Manitoba.
Canadian Field-Naturalist, 98, 2, 184–90.
Davies, S. (1984) Nomadism as a response to desert conditions in Australia. Journal of Arid Environments, 7, 183–95.
Davis, M.B. and Shaw, R.G. (2001) Range shifts and adaptive responses to quaternary climate change. Science,
292, 673–79.
Dawson, A. (2008) Control of the annual cycle in birds:
endocrine constraints and plasticity in response to ecological variability. Philosophical Transactions of the
Royal Society of London, Series B, 363, 1621–33.
De Boer, W.F. and Prins, H.H.T. (1989) Decisions of cattle
herdsmen in Burkina Faso and optimal foraging models. Human Ecology, 17, 445–64.
de Bruijn, M. and van Djik, H. (1995) Arid Ways: Cultural
Understandings of Insecurity in Fulbe Society, Central
Mali, Thela Publishers: Amsterdam.
de Leaniz, C.G. (2008) Weir removal in salmonid streams:
implications, challenges and practicalities. Hydrobiologia, 609, 83–96.
Dean, W.R.J. (2004) Nomadic Desert Birds, Springer
Verlag, London.
Dean, W.R.J., Barnard, P., and Anderson, M.D. (2009)
When to stay, when to go: trade-offs for southern African
arid-zone birds in times of drought. South African
Journal of Science, 105, 24–28.
Deinum, B. (1984) Chemical composition and nutritive
value of herbage in relation to climate. In H. Riley, and
A.O. Skjelvag (eds) The Impact of Climate on Grass
Production and Quality. Proceedings of the 10th General
Meeting of the European Grassland Federation, As,
Norway, pp. 338–50.
Del Hoyo, J., Elliott, A., and Sargatal, J. (1994) Handbook
of the Birds of the World. Volume 2: New World Vultures
to Guineafowl, Lynx Edicions: Barcelona.
D’Eon, R.G. and Delparte, D. (2005) Effects of radio-collar
position and orientation on GPS radio-collar performance, and the implications of PDOP in data screening.
Journal of Applied Ecology, 42, 383–88.
Deppe, J.L. and Rotenberry J.T. (2008) Scale-dependent
habitat use by fall migratory birds: vegetation structure,
floristics, and geography. Ecological Monographs, 78,
461–487.
Déregnacourt, S., Guyomarc’h, J., and Belhamra, M. (2005)
Comparison of migratory tendency in European quail
Coturnix c. coturnix domestic Japanese quail Coturnix
c. japonnica and their hybrids. Ibis, 147, 25–36.
Dierschke, V., Delingat, J., and Schmaljohann, H. (2003)
Time allocation in migrating northern wheatears
(Oenanthe oenanthe) during stopover: is refuelling limited by food availability or metabolically? Journal für
Ornithologie, 144, 33–44.
Dierschke, V., Mendel, B., and Schmaljohann, H. (2005)
Differential timing of spring migration in northern
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
wheatears Oenanthe oenanthe: hurried males or weak
females? Behavioral Ecology and Sociobiology, 57,
470–80.
Dingle, H. (1980) Ecology and evolution of migration. In
S.A. Gauthreaux (ed.) Animal Migration, Orientation,
and Navigation, pp. 1–100, Academic Press: London.
Dingle, H. (1989) The evolution and significance of migratory flight. In G.J. Goldsworthy (ed.) Insect Flight,
pp. 99–114, CRC Press: Boca Raton.
Dingle, H. (1996) Migration: the Biology of Life on the
Move, Oxford University Press: New York.
Dingle, H. (2006) Animal migration: is there a common
migratory syndrome? Journal of Ornithology, 147,
212–20.
Dingle, H. and Drake, V.A. (2007) What is migration?
Bioscience, 57, 113–21.
Dingle, H., Rochester, W.A., and Zalucki, M.P. (2000)
Relationships among climate, latitude and migration:
Australian butterflies are not temperate-zone birds.
Oecologia, 124, 196–207.
Dixon, J.D., Oli, M.K., Wooten, M.C., Eason, T.H., McCown,
J.W., and Paetkau, D. (2006) Effectiveness of a regional
corridor in connecting two Florida black bear populations. Conservation Biology, 20, 155–62.
Dodson, J.J. (1997) Fish migration: an evolutionary perspective. In J.-G. Godin (ed.) Behavioural Ecology of Teleost
Fishes, pp 10–36, Oxford University Press: Oxford.
Doebeli, M. (1995) Dispersal and dynamics. Theoretical
Population Biology, 47, 82–106.
Doebeli, M. and Ruxton, G. (1997) Evolution of dispersal
rates in metapopulation models: branching and cyclic
dynamics in phenotype space. Evolution, 51, 1730–41.
Dolman, P.M. and Sutherland, W.J. (1995) The response of
bird populations to habitat loss. Ibis, 137, s1, S38–S46.
Dorst, J. (1961) The Migrations of Birds, Heinemann:
London.
Dragesund, O., Johannessen, A., and Ulltang, Ø. (1997)
Variation in migration and abundance of Norwegian
spring spawning herring (Clupea harengus L.). Sarsia, 82,
2, 97–105.
Drake, V.A. and Gatehouse, A.G. (eds) (1995) Insect
Migration: Tracking Resources Through Space and
Time, Cambridge University Press: Cambridge.
Drent, R.J., Fox, A.D., and Stahl, J. (2006) Travelling to
breed. Journal of Ornithology, 147, 122–34.
Dudley, R. and Srygley, R.B. (2008) Airspeed adjustment
and lipid reserves in migratory Neotropical butterflies.
Func. Ecol., 22, 264–70.
Dudley, R., Srygley, R.B., Oliveira, E.G., and DeVries, P.J.
(2002) Flight speeds, lipid reserves, and predation of the
migratory neotropical moth Urania fulgens (Uraniidae).
Biotropica, 34, 452–58.
225
Dugger, K.M., Faaborg, J., Arendt, W.J., and Hobson, K.A.
(2004) Understanding survival and abundance of overwintering warblers: does rainfall matter? The Condor,
106, 744–60.
Duke, D.L. (2001) Wildlife use of corridors in the Central
Canadian Rockies: multivariate use of habitat characteristics and trends in corridor use. Thesis, University of
Alberta: Edmonton, Alberta, Canada.
Dulvy, N.K., Sadovy, Y., and Reynolds, J.D. (2003)
Extinction vulnerability in marine populations. Fish
and Fisheries, 4, 25–64.
Durant, S.M., Caro, T.N., Collins, D.A., Alawi, R.M. and
Fitzgibbon, C.D. (1988) Migration patterns of Thomsons
gazelles and cheetahs on the Serengeti Plains. African
Journal of Ecology, 26, 257–68.
Duriez, O., Bauer, S., Destin, A., Madsen, J., Nolet, B.A.,
Stillman, R.A., and Klaassen, M. (2009) What decision
rules might pink-footed geese use to depart on migration? An individual-based model. Behavioral Ecology,
20, 560–69.
Dusenberry, D.B. (2001) Physical constraints in sensory
ecology. In F.G. Barth and A. Schmid. (eds) Ecology of
Sensing, pp. 1–17, Springer: Berlin.
Dwyer, M. and Istomin, K. (2008) Theories of nomadic
movement: a new theoretical approach for understanding the movement decisions of Nenets and Komi reindeer herders. Human Ecology, 17, 445–64.
Ebbesson, L.O.E., Ebbesson, S.O.E., Nilsen, T.O.,
Stefansson, S.O., and Holmqvist, B. (2007) Exposure to
continuous light disrupts retinal innervation of the preoptic nucleus during parr-smolt transformation in
Atlantic salmon. Aquaculture, 273, 345–49.
Edwards, A.M. (2008) Using likelihood to test for Lévy flight
search patterns and for general power-law distributions
in nature. Journal of Animal Ecology, 77, 1212–22.
Edwards, A.M., R.A. Phillips, N.W. Watkins, M.P. Freeman,
E.J. Murphy, V. Afanasyev, S.V. Buldyrev, M.G.E. da Luz,
E.P. Raposo, H.E. Stanley, and G.M. Viswanathan. 2007.
Revisiting Levy flight search patterns of wandering albatrosses, bumblebees and deer. Nature 449:1044 –1048.
Egevang, C., Stenhouse, I.J., Phillips, R.A., Petersen,
A., Fox, J.W., and Silk, J.R.D. (2010) Tracking of Arctic
terns Sterna paradisaea reveals longest animal migration.
Proc. Natl. Acad. Sci., 107, 2078–2081, doi/10.1073/
pnas.0909493107
Eichorn, G., Drent, R.H., Stahl, J., Leito, A., and Alerstam,
T. (2009) Skippig the Baltic: the emergence of a dichotomy of alternative spring migration strategies in Russian
barnacle geese. Journal of Animal Ecology, 78, 63–72.
Ellegren, H. (1991) Stopover ecology of autumn migrating
bluethroats Luscinia svecica svecica in relation to age and
sex. Ornis Scandinavica, 22, 340–48.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
226
REFERENCES
Elliot, J.J. and Arbib, R.S.J. (1953) Origin and status of the
house finch in the eastern United States, Auk, 70,
31–37.
Ellis, D.H., Sladen, W.J.L., Lishman, W.A., Clegg, K.R.,
Duff, J.W., Gee, G.F., and Lewis, J.C. (2003) Motorized
migrations: the future or mere fantasy? BioScience, 53,
3, 260–64.
Ellis, J.E. and Swift, D.M. (1988) Stability in African pastoral ecosystems: alternate paradigms and implications
for development. Journal of Range Management, 41,
450–59.
Enloe, J.G. and David, F. (1995) Rangifer herd behaviour:
Seasonality of hunting in the Magdalenian of the Paris
Basin. In L.J. Jackson and P. T. Thacker (eds) Caribou
and Reindeer Hunters of the Northern Hemisphere,
pp. 47–63. Avebury Press: Aldershot.
Erdenebaatar, B. (2003) Mongolia case study 1: Studies on
long-distance transhumant grazing systems in Uvs and
Khuvsgul aimags of Mongolia, 1999–2000. In J.M. Suttie
and S.G. Reynolds (eds) Transhumant Grazing Systems
in Temperate Asia, pp. 31–68, Food and Agriculture
Organization of the United Nations (FAO): Rome.
Erni, B., Liechti, F., and Bruderer, B. (2002) Stopover strategies in passerine bird migration: A simulation study.
Journal of Theoretical Biology, 219, 479–93.
Ervin, J. (2003) WWF: Rapid Assessment and Prioritization
of Protected Area Management (RAPPAM) Methodology,
WWF: Gland, Switzerland.
Evans, P.R. (1966) Migration and orientation of passerine
night migrants in northeast England. Journal of Zoology,
150, 319–69.
Farnsworth, K.D. and Beecham, J.A. (1997) Beyond the
ideal free distribution: more general models of predator
distribution. Journal of Theoretical Biology, 187,
389–96.
Faugère, O., Dockes, A.C., Perrot, C., and Faugère, B. (1990a)
L’élevage traditionnel des petits ruminants au Senegal. I.
Pratiques de conduite et d’exploitation des animaux chez
les éleveurs de la région de Kolda. Revue elevage médicine vétérinaire pays tropicaux, 43, 2, 249–59.
Fedak, M., Lovell, P., McConnell, B., and Hunter, C. (2002)
Overcoming the constraints of long range radio telemetry
from animals: getting more useful data from smaller packages. Integrative and Comparative Biology, 42, 3–10.
Fedorovich, B.A. (1973) Natural conditions of the arid
zones of the USSR and ways of development of their
livestock sectors. In Essays on the Agricultural History
of the Peoples of Central Asia and Kazakhstan,
pp. 207–22, Nauka: Leningrad.
Ferguson, M.M. and Duckworth, G.A. (1997) The status
and distribution of lake sturgeon, Acipenser fulvescens, in
the Canadian provinces of Manitoba, Ontario and
Quebec: a genetic perspective. Environmental Biology
of Fishes, 48, 299–309.
Ferguson, S.H. and Elkie, P.C. (2004) Seasonal movement
patterns of woodland caribou (Rangifer tarandus caribou). Journal of Zoology, 262, 125–34.
Fernandez-Gimenez, M.E. (1997) Landscapes, Livestock,
and Livelihoods: Social, Ecological, and Land-Use
Change among the Nomadic Pastoralists of Mongolia,
University of California: Berkeley, Berkeley, CA.
Fernandez-Gimenez, M.E. (1999) Sustaining the steppes: a
geographical history of pastoral land use in Mongolia.
The Geographical Review, 89, 315–42.
Fernandez-Gimenez, M.E. (2002) Spatial and social boundaries and the paradox of pastoral land tenure: a case
study from postsocialist Mongolia. Human Ecology, 30,
49–78.
Fernandez-Gimenez, M.E. and Allen-Diaz, B. (2001)
Vegetation change along gradients from water sources
in three grazed Mongolian ecosystems. Plant Ecology,
157, 101–18.
Fernandez-Gimenez, M.E. and Batbuyan, B. (2004) Law
and disorder: Local implementation of Mongolia’s Land
Law. Development and Change, 35, 141–65.
Fernandez-Gimenez, M.E. and Le Febre, S. (2006) Mobility
in pastoral systems: dynamic flux or downward trend?
International Journal of Sustainable Development and
World Ecology, 13, 1–22.
Fernandez-Gimenez, M.E., Batbuyan, B., and Oyungerel,
J. (2007) Climate, economy and land policy: effects on
pastoral mobility patterns in Mongolia. In X. Sun and
N. Naito (eds) Mobility, Flexibility, and Potential of
Nomadic Pastoralism in Eurasia and East Africa,
pp. 3–24, Graduate School of Asian and African Area
Studies: Kyoto University, Kyoto, Japan.
Fernandez-Gimenez, M.E., Kamimura, A., and Batbuyan,
B. (2008) Implementing Mongolia’s Land Law: Progress
and Issues, Final report to the Central Asian Legal
Exchange, Central Asian Legal Exchange: Nagoya
University: Nagoya, Japan.
Ferraroli, S., Georges, J., Gaspar, P., and Le Maho, Y. (2004)
Where leatherback turtles meet fisheries. Nature, 429,
521–22.
Fieberg, J. and Delgiudice, G. (2008) Exploring migration
data using interval-censored time-to-event models.
Journal of Wildlife Management, 72, 1211–19.
Fiedler, W. (2003) Recent changes in migratory behaviour of
birds: a compilation of field observations and ringing
data. In P. Berthold, E. Gwinner, and E. Sonnenschein
(eds) Avian Migration, pp. 21–38, Springer Verlag: Berlin.
Finke, P. (2000) Changing property rights systems in
Mongolia, Max Planck Institute for Social Anthropology,
Working Paper No. 3: Halle, Germany.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Finstad G.L., Kielland K.K., and Schneider, W.S. (2006)
Reindeer herding in transition: historical and modern
day challenges for Alaskan reindeer herders. In
F. Stammler, and H. Beach (eds) People and Reindeer on
the Move, Special Issue of the journal Nomadic Peoples,
10, 2, 31–49, Oxford: Berghahn Publishers.
Fischer, J. and Lindenmayer, D.B. (2000) An assessment of
the published results of animal relocations. Biological
Conservation, 96, 1–11.
Fisher, M.J., Rao, I.M., Thomas, R.J., and Lascano, C.E.
(1996) Grasslands in the well-watered tropical lowlands.
In J.Hodgson and A.W. Illius (eds.) The Ecology and
Management of Grazing Systems, pp. 393–428, CAB
International: Wallingford, UK.
Fleishman, E., Murphy, D.D., and Brussard, P.F. (2000) A
new method for selection of umbrella species for conservation planning. Ecological Applications, 10, 2,
569–79.
Fleming, T. H. and Eby, P. (2003) Ecology of bat migration.
In T.H. Kunz and M.B. Fenton (eds) Bat Ecology,
pp. 156–208, Chicago University Press: Chicago.
Forbes, B. and Stammler, F. (2009) Arctic climate change
discourse: the contrasting politics of research agendas
in the West and Russia. Polar Research. Vol. 28, Climate
Change Vulnerability and Adaptation in the Arctic,
pp. 28–42.
Fox, A.D. and Madsen, J. (1997) Behavioural and distributional effects of hunting disturbance on waterbirds in
Europe: implications for refuge design. The Journal of
Applied Ecology, 34, 1, 1–13.
Frank, D.A., Inouye, R.S., Huntly, N., Minshall, G.W., and
Anderson J.E. (1994) The biogeochemistry of a northtemperate grassland with native ungulates—nitrogen
dynamics in Yellowstone National Park. Biogeochemistry,
26, 163–88.
Fransson, T. (1998) Patterns of migratory fuelling in
whitethroats Sylvia communis in relation to departure.
Journal of Avian Biology, 29, 569–73.
Fransson, T. and Weber, T. P. (1997) Migratory fuelling
blackcaps (Sylvia atricapilla) under perceived risk of
predation. Behavioral Ecology and Sociobiology, 41,
75–80.
Fransson, T., Barboutis, C., Mellroth, R., and Akriotis, T.
(2008) When and where to fuel before crossing the
Sahara desert—extended stopover and migratory fuelling in first-year garden warblers Sylvia borin. Journal of
Avian Biology, 39, 133–38.
Fransson, T., Jakobsson, S., Johansson, P., Kullberg, C.,
Lind, J., and Vallin, A. (2001) Bird migration—Magnetic
cues trigger extensive refuelling. Nature, 414, 35–36.
Freitag-Ronaldson, S. and Foxcroft, L.C. (2003)
Anthropogenic influences at the ecosystem level. In
227
J. Du Toit, K.H. Rogers, and H.C. Biggs (eds) The Kruger
Experience, pp. 391–421, Island Press: Washington.
Fretwell, D.D. and Lucas, H.L. 1970. On territorial behaviour and other factors influencing habitat distribution in
birds. Acta Biotheoretica, 19, 16–36.
Frey, K.E. and Smith, L.C. (2003) Recent temperature and
precipitation increases in West Siberia and their association with the Arctic Oscillation. Polar Research, 22,
287–300.
Fry, M. (2004) The status of the Saiga antelope in the
Ustiurt region of western Kazakhstan. MSc thesis,
Imperial College: London. [Online], Available: http://
www.iccs.org.uk/thesis.htm [April 2009].
Fryxell, J.M. (1991) Forage quality and aggregation by
large herbivores, The American Naturalist, 138, 478–98.
Fryxell, J.M. (1995) Aggregation and migration by grazing
ungulates in relation to resources and predators. In
A.R.E. Sinclair and P. Arcese (eds) Serengeti II: Dynamics,
Management, and Conservation of an Ecosystem, pp.
257–73, Chicago University Press: Chicago.
Fryxell, J.M. and Sinclair, A.R.E. (1988a) Seasonal migration by white-eared kob in relation to resources. African
Journal of Ecology, 26, 17–31.
Fryxell, J.M. and Sinclair, A.R.E. (1988b) Causes and consequences of migration by large herbivores. Trends in
Ecology and Evolution, 3, 237–41.
Fryxell, J. Greever, J., and Sinclair, A.R.E. (1988) Why are
migratory ungulates so abundant? American Naturalist,
131, 781–98.
Fryxell, J.M., Wilmshurst, J.F., and Sinclair, A.R.E. (2004)
Predictive models of movement by Serengeti grazers.
Ecology, 85, 2429–35.
Fryxell, J.M., Wilmshurst, J.F., Sinclair, A.R.E., Haydon,
D.T., Holt, R.D., and Abrams, P.A. (2005) Landscape
scale, heterogeneity, and the viability of Serengeti grazers. Ecology Letters, 8, 328–35.
Fryxell, J.M.,M. Hazell, L. Borger, B.D. Dalziel, D.T.
Haydon, J.M. Morales, T. McIntosh, and R.C. Rosatte.
(2008) Multiple movement modes by large herbivores at
multiple spatiotemporal scales. Proceedings of the
National Academy of Sciences of the United States of
America, 105, 19114–19.
Gagliardo, A., Ioale, P., Savini, M., and Wild, M. (2008)
Navigational abilities of homing pigeons deprived of olfactory or trigeminally mediated magnetic information when
young. Journal of Experimental Biology, 211, 2046–51.
Gallais, J. (1975) Paysans et pasteurs du Gourma. La condition sahélienne, CNRS: Paris.
Galvin, K.A., Reid, R.S., Behnke, R.H., and Hobbs, N.T.
(eds) (2008) Fragmentation in Semi-Arid and Arid
Landscapes: Consequences of Human and Natural
Systems, Springer: Dordrecht, The Netherlands.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
228
REFERENCES
Gannes, L.Z. (2002) Mass change pattern of blackcaps
refueling during spring migration: Evidence for physiological limitations to food assimilation. Condor, 104,
231–39.
Garin, P., Faye, A. Lericollais, A., and Sissokho, M. (1990)
Évolution du rôle du bétail dans la gestion de la fertilité
des terroirs Sereer au Sénégal. Les Cahiers de la
Recherche Développement, 26, 65–84.
Gatehouse, A.G. (1987) Migration: a behavioral process
with ecological consequences? Antenna, 11, 10–12.
Gauthreaux, S. (1982) The ecology and evolution of avian
migration systems. Avian Biology, 6, 93–168.
Gaylard, A., Owen-Smith, N., and Redfern J. (2003) Surface
water availability: implications for heterogeneity and
ecosystem processes. In J. Du Toit, K. H. Rogers, and
H. C. Biggs (eds) The Kruger Experience, pp. 171–88,
Island Press: Washington.
Geist, V. (1999) Deer of the World: Their Evolution,
Behaviour, and Ecology, Swan Hill Press: Shrewsbury.
Geller, M. K. and Vorzhonov, V. V. (1975) Migratsii i sezonnoe razmeshchenie dikikh severnykh olenei taimyrskoi
populiatsii. In E. E. Syroechkovski (ed.) Dikii Severnyi
Olen’ v SSSR, pp. 80–88, Central Board of Game
Management and Nature Reserves of the RSFSR:
Moscow.
Gelman, A. and J. Hill (2007) Data Analysis using Regression
and Multilevel/Hierarchical Models, Cambridge University Press: Cambridge.
Georgiadis, N.J. and McNaughton, S.J. (1990) Elemental
and fibre contents of savanna grasses: variation with
grazing, soil type, season and species. Journal of Applied
Ecology, 27, 623–34.
Gerber, L.R., Heppell, S.S., Ballantyne, F., and Sala, E.
(2005) The role of dispersal and demography in determining the efficacy of marine reserves. Canadian Journal
of Fisheries and Aquatic Sciences, 62, 863–71.
Gereta, E., Mwangomo, E., and Wolanski, E. (2009)
Ecohydrology as a tool for the survival of the threatened
Serengeti ecosystem. Ecohydrology and hydrobiology,
9, 115–24.
Gerlier, M. and Roche, P. (1998) A radio telemetry study of
the migration of Atlantic salmon (Salmo salar L.) and sea
trout (Salmo trutta trutta L.) in the upper Rhine.
Hydrobiologia, 371/372, 283–93.
Gibo, D.L. and Pallett, M.J. (1979) Soaring flight of monarch butterflies, Danaus plexippus (Lepidoptera, Danaidae), during the late summer migration in Southern
Ontario. Canadian Journal of Zoology, 57, 1393–1401.
Gilbert, M., Slingenbergh, J., and Xiao, X. (2008) Climate
change and avian influenza. Revue Scientifique Et
Technique-Office International Des Epizooties, 27,
459–66.
Gill, R.E., Tibbitts, T.L., Douglas, D.C., Handel, C.M.,
Mulcahy, D.M., Gottschalck, J.C., Warnock, N.,
McCaffery, B.J., Battley, P.F., and Piersma T. (2009)
Extreme endurance flights by landbirds crossing the
Pacific Ocean: ecological corridor rather than barrier?
Proceedings of the Royal Society B-Biological Sciences,
276, 447–58.
Giller, P.S. (2005) River restoration: seeking ecological
standards. Editor’s introduction. Journal of Applied
Ecology, 42, 201–207.
Gilly, W.F., Markaida, U., Baxter, C.H., et al. (2006) Vertical
and horizontal migrations by the jumbo squid Dosidicus
gigas revealed by electronic tagging. Marine EcologyProgress Series, 324, 1–17.
Gilyazov, A. and Sparks, T.H. (2002) Change in the timing
of migration of common birds at the Lapland Nature
Reserve (Kola Peninsula, Russia) during 1931–1999.
Avian Ecology and Behaviour, 8, 35–47.
Gloyne, C.C. and Clevenger, A.P. (2001) Cougar (Puma
concolor) use of wildlife crossing structures on the TransCanada highway in Banff National Park, Alberta.
Wildlife Biology, 7, 2, 117–24.
Godley, B.J., Broderick, A.C., and Hays, G.C. (2001) Nesting
of green turtles (Chelonia mydas) at Ascension Island,
South Atlantic. Biological Conservation, 97, 151–58.
Godley, B.J., Lima, E.H.S.M., Åkesson, S., Broderick, A. C.,
Glen, F., Godfrey, M.H., Luschi, P. and Hays, G.C. (2003)
Movement patterns of green turtles in Brazilian coastal
waters described by satellite tracking and flipper tagging. Marine Ecology Progress Series, 253, 279–88.
Goldstein, D.L. and Pinshow, B. (2006) Taking physiology
to the field: using physiological approaches to answer
questions about animals in their environments.
Physiological and Biochemical Zoology, 79, 237–41.
Goodchild, M.F., Friedl, M.A., and Case, T.J. (eds)
(2001) Spatial Uncertainty in Ecology, Springer-Verlag:
New-York,
Goodhue, R.E. and McCarthy, N. (1999) Fuzzy access:
modelling grazing rights in sub-Saharan Africa. In
N. McCarthy, D. Swallow, M. Kirk and P. Hazell (eds)
Property Rights, Risk, and Livestock Development in
Africa, pp. 191–210, International Food Policy Research
Institute: Washington, DC.
Gordo, O. (2007) Why are bird migration dates shifting?
A review of weather and climate effects on avian
migratory phenology. Climate Research, 35, 37–58.
Granberg, L., Soini K., and Kantanen. J. (eds) (2009)
Sakha Ynaga—Cattle of the Yakuts. Annales Academiae
Scientiarum Fennicae Humaniora 355: Helsinki.
Submitted.
Gray, P. (2006) ‘The Last Kulak’ and other stories of postprivatization life in Chukotka’s tundra, in People and
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Reindeer on the Move. In F. Stammler, and H. Beach
(eds) Special Issue of the journal Nomadic Peoples, 10,
2, pp 50–67, Berghahn Publishers: Oxford.
Green, A.A. (1988) Elephants of the Pendjari-SingouMekrou region, West Africa. Mammalia, 52, 4, 557–65.
Green, J.A., Halsey, L.G., Wilson, R.P., and Frappell, P.B.
(2009) Estimating energy expenditure of animals using
the accelerometry technique: activity, inactivity and
comparison with the heart-rate technique. Journal of
Experimental Biology, 212, 471–82.
Green, M. and Alerstam, T. (2000) Flight speeds and climb
rates of Brent geese: mass-dependent differences
between spring and autumn migration. Journal of Avian
Biology, 31, 215–25.
Green, M., Alerstam, T., Clausen, P., Drent, R., and Ebbinge,
R.S. (2002) Dark-bellied Brent geese Branta bernicla bernicla, as recorded by satellite telemetry, do not minimize
flight distance during spring migration. Ibis, 144, 106–21.
Green, M., Alerstam, T., Gudmundsson, G.A., Hedenstrom,
A., and Piersma, T. (2004) Do Arctic waders use adaptive wind drift? Journal of Avian Biology, 35, 305–15.
Greenwood, P.J. (1980) Mating systems, philopatry, and
dispersal in birds and mammals. Animal Behavior, 28,
1140–62.
Griffioen, P.A. and Clarke, M.F. (2002) Large-scale birdmovement patterns evident in eastern Australian atlas
data. Emu, 102, 97–125.
Griffith, B., Scott, J.M., Carpenter, J.W., and Reed C. (1989)
Translocation as a species conservation tool: status and
strategy. Science, 245, 477–82.
Grimm, V.,E. Revilla, U. Berger, F. Jeltsch, W.M. Mooij,
S. F. Railsback, H.H. Thulke, J. Weiner, T. Wiegand, and
D.L. DeAngelis (2005) Pattern-oriented modeling of
agent-based complex systems: lessons from ecology.
Science, 310, 987–91.
Grinnel, J. (1931) Some angles in the problem of bird
migration. Auk, 48, 22–32.
Griswold, C.K., Taylor, C.M., and Norris D.R. (2010) The
evolution of migration in a seasonal environment.
Proceedings of the Royal Society of London (B),
doi:10.1.1098/rspb.2010.0550.
Gross, M.R. (1987) Evolution of diadromy in fishes.
American Fisheries Society Symposium, 1, 14–25.
Grouzis, M. (1988) Structure, productivité, et dynamique
des systèmes écologiques Sahéliens (Mare d’Oursi,
Burkina Faso), Collection Études et Thèses, Éditions de
l’ORSTOM: Paris.
Grunbaum, D. (1998a) Schooling as a strategy for taxis in a
noisy environment. Evolutionary Ecology, 12, 503–22.
Grunbaum, D. (1998b) Using spatially explicit models to
characterize foraging performance in heterogeneous
landscapes. American Naturalist, 151, 97–115.
229
Gschweng, M., Kalko, E.K.V., Querner, U., Fiedler, W.,
and Berthold, P. (2008) All across Africa: highly
individual migration routes of Eleonora’s falcon.
Proceedings of the Royal Society B-Biological Sciences,
275, 2887–96.
Gudmundsson, G.A. and Alerstam, T. (1998) Optimal map
projections for analysing long-distance migration
routes. J Avian Biol., 29, 597–605.
Gudmundsson, G.A., Linddström, Å., and Alerstam, T.
(1991) Optimal fat loads and long-distance flights by
migrating knots Calidris canutus, sanderling C. alba and
turnstones Arenaria interpres. Ibis, 133, 140–52.
Guillemain, M., Bertout, J.M., Christensen, T.K., Poysa,
H., Vaananen, V.M., Triplet, P., Schricke V., et al. (2010)
How many juvenile Teal Anas crecca reach the wintering
grounds? Flyway-scale survival rate inferred from wing
age-ratios. Journal of Ornithology, 151, 51–60.
Gulliver, P.H. (1975) Nomadic movements: causes and
implications. In T. Monod (ed.) Pastoralism in Tropical
Africa, pp. 369–86, OUP: London.
Gunderson, H., Andreassen, H.P., and Storaas, T. (2004)
Supplemental feeding of migratory moose Alces alces:
forest damage at two spatial scales. Wildlife Biology, 10,
213–23.
Gunn, A. (1998) Caribou and muskox harvesting in the
Northwest Territories. In E.J. Milner-Gulland and R.
Mace (eds) Conservation of Biological Resources, pp.
314–30, Blackwell Science: Oxford.
Gustafson, T., Lindkvist, B., Gotborn, L., and Gyllin, R.
(1977) Altitudes and flight times for swifts Apus apus L.
Ornis Scandinavica, 8, 87–95.
Gustafson, R.G., Waples, R.S., Myers, J.M., Weitkamp,
L.A., Bryant, G.J., Johnson, O.W., and Hard, J.J. (2007)
Pacific salmon extinctions: quantifying lost and remaining diversity. Conservation Biology, 21, 4, 1009–20.
Gwinner, E. (1969) Untersuchungen zur Jahresperiodik
von Laubsängern. Die Entwicklung des Gefieders, des
Gewichts und der Zugunruhe bei Jungvögeln der Arten
Phylloscopus bonelli, Ph. sibilatrix und Ph. collybita. Journal
of Ornithology, 110, 1–21.
Gwinner, E. (1977) Circannual rhythms in bird migration.
Annual Review of Ecology and Systematics, 8,
381–405.
Gwinner, E. (1990) Bird Migration: Physiology and
Ecophysiology. Springer-Verlag: Berlin.
Gwinner, E. (1996) Circadian and circannual programmes
in avian migration. The Journal of Experimental Biology,
199, 39–48.
Gwinner, E. and Wiltschko, W. (1978) Endogenously controlled changes in migratory direction of the Garden
Warbler, Sylvia borin. Journal of Comparative Physiology,
125, 267–73.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
230
REFERENCES
Gwinner, E. and Wiltschko, W. (1980) Circannual
changes in migratory orientation of the Garden
Warbler, Sylvia borin. Behavioral Ecology and
Sociobiology, 7, 73–78.
Habeck, J. O. (2005) What it means to be a herdsman: The
Practice and Image of Reindeer Husbandry among the
Komi of Northern Russia, Vol. 5, Halle Studies in the
Anthropology of Eurasia, Lit: Münster
Habou, A., and Danguioua, A. (1991) Transfert du capital –
Bétail au Niger (des pasteurs aux autres catégories
socio-professionnelles), Secrétariat Permanent du
Comite National du Code Rural: Niamey, Niger.
Hahn, S., Bauer, S., and Liechti F. (2009) The natural link
between Europe and Africa—2.1 billion birds on migration. Oikos, 118, 624–26.
Haining, R. (2003) Spatial Data Analysis: Theory and
Practice, Cambridge University Press: Cambridge.
Hamelin, F. and Lewis, M. (2010) A differential game theoretical analysis of mechanistic models for territoriality,
Journal of Mathematical Biology, 61, 665–694.
Hamilton, W.D. (1971) Geometry for the selfish herd.
Journal of Theoretical Ecology, 31, 295–311.
Hancock, P.A. and Hutchinson, M.F. (2006) Spatial
Interpolation of large climate data sets using thin plate
smoothing splines. Environmental Modelling and
Software, 21, 1684–94.
Hancock, P.A. and Milner-Gulland, E. J. (2006) Optimal
movement strategies for social foragers in unpredictable
environments. Ecology, 87, 2094–2102.
Hannesson, R. (2004) Sharing the Herring: Fish Migrations,
Strategic Advantage, and Climate Change, SNF Working
Paper No. 27/2004. [Online], Available: http://bora.
nhh.no:8080/bitstream/2330/382/1/A2704.pdf
[January 2009].
Hanrahan T.P., Dauble, D.D., and Geist, D.R. (2004) An
estimate of chinook salmon (Oncorhynchus tshawytscha)
spawning habitat and redd capacity upstream of a
migration barrier in the upper Columbia River. Canadian
Journal of Fisheries and Aquatic Sciences, 61, 1, 23–33.
Hanski, I. (1998) Metapopulation dynamics. Nature, 396,
41–49.
Hansson, L. A. and Hylander, S. (2009) Size-structured
risk assessments govern Daphnia migration. Proceedings
of the Royal Society B-Biological Sciences, 276, 331–36.
Haro, A., Richkus, W., Whalen, K., Hoar, A., Busch, W.-D.,
Lary, S., Brush, T., and Dixon, D. (2000) Population
decline of the American eel: implications for research
and management. Fisheries, 25, 9, 7–16.
Harrington, R., Owen-Smith, N., Viljoen, P.C., Biggs, H.C.,
Mason, D.R., and Funston, P. (1999) Establishing the
causes of the roan antelope decline in the Kruger National
Park, South Africa. Biological Conservation, 90, 69–78.
Harris, G., Thirgood, S., Grant, J., Hopcraft, C., Cromsigt,
J.P.G.M., and Berger, J. (2009) Global decline in aggregated migrations of large terrestrial mammals.
Endangered Species Research, 7, 55–76.
Harvey, P.H. and Pagel, M.D. (1991) The comparative
method in evolutionary biology, Oxford University
Press: Oxford.
Hary, I., Schwartgz, H-J., Peilert, V.H.C., and Mosler C.
(1996) Land degradation in African pastoral systems
and the destocking controversy. Ecological Modelling,
89, 227–33.
Hasegawa, H. and DeGange, A.R. (1982) The short-tailed
albatross, Diomedea albatrus, its status, distribution and
natural history. American Birds, 36, 806–14.
Haslam, C. (2008) Nobody to watch wildebeest migration
in Kenya. The Sunday Times, 8 June [Online], Available:
http://www.timesonline.co.uk/tol/travel/news/article4076148.ece. [November 2008].
Hastie, T., Tibshirani, R., and Friedman, J. (2001) The elements of statistical learning: data mining, inference, and
prediction: Springer Series in Statistics, Springer:
New York.
Hawkes, L.A., Broderick, A.C., Coyne, M.S., Godfrey,
M.H., and Godley, B.J. (2007) Only some like it hot—
quantifying the environmental niche of the loggerhead
sea turtle. Diversity and Distributions, 13, 447–57.
Hawkes, L.A., Broderick, A.C., Coyne, M.S., Godfrey, M.S.,
Lopez-Jurado, L.F., Lopez-Suarez, P., Merino, S.E., VaroCruz, N., and Godley, B.J. (2006) Phenotypically linked
dichotomy in sea turtle foraging requires multiple conservation approaches. Current Biology, 16, 990–95.
Hay, R.K.M. and Heide, O.M. (1984) The response of highlatitude Norwegian grass cultivars to long photoperiods and cool temperatures. In H. Riley and A.O Skjelvag
(eds) The Impact of Climate on Grass Production and
Quality, Proceedings of the 10th General Meeting of
the European Grassland Federation, As, Norway,
pp. 46–50.
Haydon, D.T., Morales, J.M., Yott, A., Jenkins, D.A.,
Rosatte, R., and Fryxell, J. (2008) Socially-informed random walks: Incorporating group dynamics into models
of population spread and growth. Proceedings of the
Royal Society B-Biological Sciences, 275, 1101–1109.
Hays, G.C. (2008) Sea turtles: A review of some key recent
discoveries and remaining questions. Journal of
Experimental Marine Biology and Ecology, 356, 1–7.
Hays, G.C., Houghton, J.D.R., and Myers, A.E. (2004)
Endanqered species—Pan-Atlantic leatherback turtle
movements. Nature, 429, 522.
Healey, M. C. and Groot, C. (1987) Marine migration and
orientation of ocean-type Chinook and sockeye salmon.
American Fish Society Symposium, 1, 298–312.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Heath, M.R., Boyle, P.R., Gislason, A., Gurney, W.S.C.,
Hay, S.J., Head, E.J.H., Holmes, S., Ingvarsdottir, A.,
Jonasdottir, S.H., Lindeque, P., Pollard, R.T., Rasmussen,
J., Richards, K., Richardson, K., Smerdon, G. and Speirs,
D. (2004) Comparative ecology of over-wintering
Calanus finmarchicus in the northern North Atlantic,
and implications for life-cycle patterns. ICES Journal of
Marine Science, 61, 698–708.
Hebblewhite, M. and Merrill, E. (2007) Multiscale wolf
predation risk for elk: does migration reduce risk?
Oecologia, 152, 377–87.
Hebblewhite, W. and Merrill, E.H. (2009) Trade-offs
between predation risk and forage differ between
migrant strategies in a migratory ungulate. Ecology, 90,
3445–54.
Hebblewhite, M., Merrill, E. and McDermid, G. (2008) A
multi-scale test of the forage maturation hypothesis in a
partially migratory ungulate population. Ecological
Monographs, 78, 141–66.
Hebblewhite, M., White, C.A., Nietvelt, C.G., McKenzie,
J.A., Hurd, T.E., Fryxell, J.M., Bayley, S.E. and Paquet, P.
(2005) Human activity mediates a trophic cascade
caused by wolves. Ecology, 86, 2135–44.
Hedenström, A. (1992) Flight performance in relation to
fuel load in birds. Journal of Theoretical Biology, 158,
535–37.
Hedenström, A. (1993) Migration by soaring or flapping flight in birds: the relative importance of energy
cost and speed. Philosophical Transactions of the
Royal Society of London B Biological Sciences, 342,
353–61.
Hedenström, A. (2001) Twenty-three testable predictions
about bird flight. In P. Berthold, E. Gwinner, and E.
Sonnenschein (eds) Avian Migration, pp. 563–82,
Springer-Verlag: Berlin.
Hedenström, A. (2003a) Optimal migration strategies in
animals that run: a range equation and its consequences.
Animal Behaviour, 66, 631–36.
Hedenström, A. (2003b) Scaling migration speed in animals that run, swim and fly. J. Zool., 259, 155–60.
Hedenström, A. (2006) Scaling of migration and the annual
cycle of birds. Ardea, 94, 399–408.
Hedenström, A. (2008) Adaptations to migration in
birds: behavioural strategies, morphology and scaling
effects. Philosophical Transactions of the Royal Society
B, 363, 287–99. doi:10.1098/rstb.2007.2140.
Hedenström, A. (2009) Optimal migration strategies in
bats. Journal of Mammalogy, 90, 1298–1309.
Hedenström, A. and Alerstam, T. (1992) Climbing performance of migrating birds as a basis for estimating
limits for fuel-carrying capacity and muscle work.
Journal of Experimental Biology, 164, 19–38.
231
Hedenström, A. and Alerstam, T. (1996) Skylark optimal
flight speeds for flying nowhere and somewhere.
Behavioral Ecology, 7, 121–26.
Hedenström, A. and Alerstam, T. (1997) Optimum fuel
loads in migratory birds: distinguishing between time
and energy minimization. Journal of Theoretical Biology,
189, 227–34.
Hedenström, A. and Pettersson, J. (1986) Differences in fat
deposits and wing pointedness between male and
female willow warblers caught on spring migration at
Ottenby, SE Sweden. Ornis Scandinavica, 17, 182–85.
Hedenström, A. and Pettersson, J. (1987) Migration routes
and wintering areas of willow warblers Phylloscopus
trochilus (L) ringed in Fennoscandia. Ornis Fennica, 64,
137–43.
Hedenström, A., Johansson, L.C., and Spedding, G.R.
(2009) Bird or bat: comparing airframe design and flight
performance. Bioinspiration and Biomimetics, 4, 1.
Hedenström, A., Barta, Z., Helm, B., Houston, A.I.,
McNamara, J.M., and Jonzén, N. (2007) Migration speed
and scheduling of annual events by migrating birds in
relation to climate change. Climate Research, 35, 79–91.
Heino, M. and Hanski, I. (2001) Evolution of migration
rate in a spatially realistic metapopulation model.
American Naturalist, 157, 495–511.
Helbig, A. J. (1991) Inheritance of migratory direction in a
bird species—A cross-breeding experiment with
SE-migrating and SW-migrating blackcaps (Sylvia
atricapilla). Behavioral Ecology and Sociobiology, 28, 9–12.
Helbig, A.J. (2003) Evolution of migration: a phylogenetic
and biogeographic perspective. In P Berthold, E
Gwinner and E Sonnenschein (eds) Avian Migration,
pp. 3–20. Springer-Verlag: Heidelberg.
Helfield, J.M. and Naiman R.J. (2001) Effects of
salmon-derived nitrogen on riparian forest growth
and implications for stream productivity. Ecology,
82, 2403–2409.
Helm, B., Gwinner, E., and Trost, L. (2005) Flexible seasonal timing and migratory behavior: results from
Stonechat breeding programs. Annals of the New York
Academy of Sciences, 1046, 216–27.
Helms, C.W. (1963) The annual cycle and zugunruhe in
birds. Proceedings of the International Ornithological
Congress, 13, 925–39.
Hiernaux, P. (1998) Effects of grazing on plant species
composition and spatial distribution in rangelands of
the Sahel. Plant Ecology, 138, 2, 191–202.
Hiernaux, P. and L. Diarra. (1984) Savanna burning, a controversial technique for rangeland management in the Niger
flood plains of central Mali. In J.C. Tothill and J.J. Mott
(eds) Ecology and Management of the World’s Savannas,
pp. 238–43, Australian Academy of Sciences: Canberra.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
232
REFERENCES
Hiernaux, P. and Turner, M.D. (1996) The effect of the timing and frequency of clipping on nutrient uptake and
production of Sahelian annual rangelands. Journal of
Applied Ecology, 33, 387–99.
Hilborn, R. and Mangel, M. (1997) The Ecological
Detective: Confronting Models with Data. Monographs
in Population Biology, Princeton University Press:
Princeton.
Hind, A., Gurney, W.S.C., Heath, M., and Bryant, A.D.
(2000) Overwintering strategies in Calanus finmarchicus.
Marine Ecology Progress Series, 193, 95–107.
Hirche, H.J. (1996) Diapause in the marine copepod,
Calanus finmarchicus—a review. Ophelia, 44, 129–43.
Hjeljord, O. (2001) Dispersal and migration in northern
forest deer—are there unifying concepts? Alces, 37, 2,
353–70.
Hoar, W. S. (1988) The physiology of smolting salmonids.
In W. S. Hoar and D. J. Randall (eds) Fish Physiology,
pp. 275–343, Academic Press: London.
Hobbs, N.T. and Swift, D.M. (1988) Grazing in herds:
when are nutritional benefits realized? American
Naturalist, 131, 760–64.
Hobson, K.A. and D.R. Norris. (2008) Animal migration: a
context for using new techniques and approaches. In
K.A. Hobson and L. I. Wassenaar (eds) Tracking Animal
Migration with Stable Isotopes, pp. 1–19 Terrestrial
Ecology Series, Elsevier: Amsterdam.
Hobson, K.A. and Wassenaar, L.I. (2008), Tracking Animal
Migration with Stable Isotopes: Terrestrial Ecology
Series, Elsevier.
Holdo, R.M. (2007) Elephants, fire, and frost can determine community structure and composition in Kalahari
woodlands. Ecological Applications, 17, 558–68.
Holdo, R.M., Holt, R.D., and Fryxell, J.M. (2009a) Grazers,
browsers, and fire influence the extent and spatial pattern of tree cover in the Serengeti. Ecological
Applications, 19, 95–109.
Holdo, R.M., Holt, R.D., and Fryxell, J.M. (2009b) Opposing
rainfall and nutrient gradients best explain the wildebeest migration in the Serengeti. The American
Naturalist, 173, 431–45.
Holdo, R.M., Holt, R. Coughenour, D.M.B., and Ritchie,
M.E. (2007) Plant productivity and soil nitrogen as a
function of grazing, migration and fire in an African
savanna. Journal of Ecology, 95, 115–28.
Holdo, R.M., Sinclair, A.R.E., Metzger, K.L., Bolker, B.M.,
Dobson, A.P., Ritchie, M.E., and Holt, R.D. (2009c) A
disease-mediated trophic cascade in the Serengeti:
implications for ecosystem structure and C stocks. PLOS
Biology, 7, e1000210.
Holland, R.A., Borissov, I., and Siemers, B.M. (2010) A nocturnal mammal, the greater mouse-eared bat, calibrates
a magnetic compass by the sun. Proceedings of the
National Academy of Sciences, 107, 15, 6941–45.
Holland, R.A., Wikelski, M., and Wilcove, D.S. (2006b)
How and why do insects migrate? Science, 313, 794–96.
Holland, R. A., Wikelski, M., Kummeth, F., and Bosque, C.
(2009) The secret life of oilbirds: new insights into the
movement ecology of a unique avian frugivore. PLoS
ONE, 4, 12, e8264. doi:10.1371/journal.pone.0008264.
Holland, R.A., Thorup, K., Vonhof, M.J., Cochran, W.W.,
and Wikelski, M. (2006a) Navigation—bat orientation
using Earth’s magnetic field. Nature, 444, 702.
Holst, J.C., Rottingen, I., and Melle, W. (2004) The herring.
In H.R. Skjoldal (ed.) The Norwegian Sea Ecosystem,
pp. 203–26, Tapir Academic Press: Trondheim.
Holt, R.D. (1997) On the evolutionary stability of sink
populations. Evolutionary Ecology, 11, 723–31.
Holt, R.D. (2004) Implications of system openness for
local community structure and ecosystem function. In
G. E. Polis, M. E. Power, and G. R. Huxel (eds) Food
Webs at the Landscape Level, pp. 96–114, University of
Chicago Press: Chicago.
Holt, R.D. (2008) Theoretical perspectives on resource
pulses. Ecology, 89, 671–81.
Holt, R.D. and McPeek, M.A. (1996) Chaotic population
dynamics favors the evolution of dispersal. American
Naturalist, 148, 709–18.
Holyoak, M., Leibold, M.A., Mouquet, N.M., Holt, R.D.,
and Hoopes M.F. (2005) Metacommunities: a framework
for large-scale community ecology. In M. Holyoak, M.A.
Leibold, and R.D. Holt (eds) Metacommunities: Spatial
Dynamics and Ecological Communities, pp. 1–32
University of Chicago Press: Chicago.
Horne, J.S., Garton, E.O., Krone, S.M., and Lewis, J.S.
(2007) Analyzing animal movements using Brownian
bridges. Ecology, 88, 2354–63.
Houston, A.I. (1998) Models of optimal avian migration:
state, time and predation. Journal of Avian Biology, 29,
395–404.
Houston, A.I. and McNamara, J.M. (1999) Models of
Adaptive Behaviour: An Approach Based on State,
Cambridge University Press: Cambridge.
Houston, A.I., Stephens, P.A., Boyd, I.L., Harding, K.C.,
and McNamara, J.M. (2007) Capital or income breeding? A theoretical model of female reproductive strategies. Behavioral Ecology, 18, 241–50.
Hume, I.D. and Biebach, H. (1996) Digestive tract function
in the long-distance migratory garden warbler, Sylvia
borin. Journal of Comparative Physiology B, 166,
388–95.
Humphrey, C. and Sneath, D. (1999) The End of
Nomadism? Society, State and the Environment in Inner
Asia, Duke University Press: Durham, NC.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Hunsaker, C.T., Goodchild, M.F., Friedl, M.A., and Case,
T.J. (2001) Spatial Uncertainty in Ecology, Springer
Verlag: New-York.
Hunter, E., Metcalfe, J.D., O’Brien, C.M., Arnold, G.P., and
Reynolds, J.D. (2004) Vertical activity patterns of freeswimming adult plaice in the southern North Sea.
Marine Ecology-Progress Series, 279, 261–73.
Hurford, A. (2009) GPS measurement error gives rise to
spurious 180° turning angles and strong directional
biases in animal movement data. PLoS ONE, 4, e5632.
Hurry, G.D., Hayashi, M., and Maguire, J.J. (2008) Report
of the independent review of ICCAT, ICCAT document
PLE-106/2008. [Online}, Available: http://www.iccat.
int/com2008/ENG/PLE-106.pdf.
Huse, G. and Giske, J. (1998) Ecology in Mare Pentium: an
individual-based spatio-temporal model for fish with
adapted behaviour. Fisheries Research, 37, 163–78.
Huse, G., Railsback, S., and Fernö, A. (2002) Modelling
changes in migration pattern of herring: collective
behaviour and numerical domination. Journal of Fish
Biology, 60, 571–82.
Huse, G., Strand, E., and Giske, J. (1999) Implementing
behaviour in individual-based models using neural networks and genetic algorithms. Evolutionary Ecology,
13, 469–83.
Hutterer, R., Ivanova, T., Meyer-Cords, C., and Rodrigues,
L. (2005) Bat migrations in Europe: a review of banding
data and literature, Federal Agency for Nature
Conservation: Bonn, Germany.
Hvenegaard, G.T., Butler, J.R., and Krystofiak, D.K. (1989)
Economic values of bird watching at Point Pelee
National Park, Canada. Wildlife Society Bulletin, 17, 4,
526–31.
ICES (The International Council for the Exploration of the
Sea) (2007) Report of the Working Group on Northern
Pelagic and Blue Whiting Fisheries, 27 August–1
September 2007, Vigo, Spain, [Online], Available:
http://www.ices.dk/reports/ACOM/2007/WGNPBW/
ACFM2907.pdf. [February 2009].
ICES (The International Council for the Exploration of the
Sea) (2008a) Norwegian spring-spawning herring.
Report of the Advisory Committee on Fisheries
Management, [Online], Available: http://www.ices.
dk/committe/acom/comwork/report/2008/2008/
her-noss.pdf. [December 2008].
ICES (The International Council for the Exploration of the
Sea) (2008b) Report of the Working Group on Widely
Distributed Stocks (WGWIDE), 2–11 September 2008,
ICES CM 2008/ACOM, 13.
Illius, A. and Gordon, I. (1992) Modeling the nutritional
ecology of ungulate herbivores: evolution of body size
and competitive interactions. Oecologia, 89, 428–34.
233
Illius A. and O’Connor, T. (1999) On the relevance of nonequilibrium concepts to semi-arid grazing systems.
Ecological Applications, 8, 798–813.
Inglis, J.M. (1976) Wet season movements of individual
wildebeests of the Serengeti migratory herd. East
African Wildlife Journal, 14, 17–34.
Ingold, T. (1980) Hunters, Pastoralists and Ranchers,
Cambridge University Press: Cambridge.
Inouye, D.W., Barr, B., Armitage, K.B., and Inouye, B.D.
(2000) Climate change is affecting altitudinal migrants
and hibernating species. Proceedings of the National
Academy of the Sciences, 97, 4, 1630–33.
IOTC (Indian Ocean Tuna Commission) (2008) Report of the
Fourth Session of the IOTC Working Party on Ecosystems
and Bycatch, 20–22 October 2008, Bangkok, Thailand.
IPCC (Intergovernmental Panel on Climate Change) (2001)
Climate Change 2001: The Scientific Basis. Contribution
of Working Group I to the Third Assessment Report of
the IPCC, J.T. Houghton, Y. Ding, D.J. Griggs, M.
Noguer, P.J. van der Linden, X. Dai, K. Maskell and C.A.
Johnson (eds) Cambridge University Press: Cambridge.
Ito, T., Miura, N., Lhagvasuren, B., Enkhbileg, D.,
Takatsuki, S., Tsunekawa, A., and Jiang, Z. (2005)
Preliminary evidence of a barrier effect of a railroad on
the migration of Mongolian gazelles. Conservation
Biology, 19, 945–48.
Ito, T.Y., Miura, N., Lhagvasuren, B., Enkhbileg, D.,
Takatsuki, S., Tsunekawa, A., and Jiang, Z. (2006)
Satellite tracking of Mongolian gazelles (Procapra gutturosa) and habitat shifts in their seasonal ranges. Journal
of Zoology, 269, 291–98.
Izhaki, I. and Safriel, U.N. (1985) Why do fleshy-fruit
plants of the Mediterranean scrub intercept fall- but not
spring-passage of seed-dispersing migratory birds?
Oecologia, 67, 40–43.
Jaeger, M.E., Zale, A.V., McMahon, T.E., and Schmitz, B.J.
(2005) Seasonal movements, habitat use, aggregation,
exploitation, and entrainment of saugers in the lower
Yellowstone River: an empirical assessment of factors
affecting population recovery. North American Journal
of Fisheries Management, 25, 1550–68.
Jamemo, A. (2008) Seasonal migration of male red deer
(Cervus elaphus) in southern Sweden and consequences
for management. European Journal of Wildlife Research,
54, 327–33.
Jefferies, R.L., Jano, A.P., and Abraham, K.F. (2006) A biotic
agent promotes large-scale catastrophic change in the
coastal marshes of Hudson Bay. Journal of Ecology, 94,
234–42.
Jenni, L. and Jenni-Eiermann, S. (1992) Metabolic patterns of
feeding, overnight fasted and flying night migrants during autumn migration. Ornis Scandinavica, 23, 251–59.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
234
REFERENCES
Jenni, L. and Jenni-Eiermann, S. (1998) Fuel supply and
metabolic constraints in migrating birds. Journal of
Avian Biology, 29, 521–28.
Jenni, L. and Kéry, M. (2003) Timing of autumn bird migration under climate change: advances in long-distance
migrants, delays in short-distance migrants. Proceedings
of the Royal Society of London Series B, 270, 1523,
1467–71.
Jenni-Eiermann, S., Jenni, L., Kvist, A., Lindström, A.,
Piersma, T., and Visser, G.H. (2002) Fuel use and metabolic response to endurance exercise: a wind tunnel
study of a long-distance migrant shorebird. Journal of
Experimental Biology, 205, 2453–60.
Jiang, Z., Takatsuki, S., Li, J., Wang, W.,Gao, Z., and Ma, J.
(2002) Seasonal variation in foods and digestion of
Mongolian gazelles in China. Journal of Wildlife
Management, 66, 1, 40–47.
Johnson, C.G. (1969) Migration and Dispersal of Insects by
Flight, Methuen: London.
Johnson, C.J., Parker, K.L., Heard, D.C. and Gillingham,
M.P. (2002) Movement parameters of ungulates and
scale-specific responses to the environment. Journal of
Animal Ecology, 71, 225–35.
Jones, J., Barg, J.J., Sillett, T.S., Veit, M.L., and Robertson,
R.J. (2004) Minimum estimates of survival and population growth for cerulean warblers (Dendroica cerulea)
breeding in Ontario, Canada. Auk, 121, 15–22.
Jonsen, I.D., Flenming, J.M., and Myers, R.A. (2005) Robust
state-space modeling of animal movement data. Ecology,
86, 2874–80.
Jonsen, I.D., Myers, R.A., and James, M.C. (2006) Robust
hierarchical state-space models reveal diel variation in
travel rates of migrating leatherback turtles. Journal of
Animal Ecology, 75, 1046–57.
Jonsson, B. and Jonsson, N. (1993) Partial migration: niche
shift versus sexual maturation in fishes. Reviews in Fish
Biology and Fisheries, 3, 4, 348–65.
Jonsson, B. and N. Jonsson (2003) Migratory Atlantic
salmon as vectors for the transfer of energy and nutrients between freshwater and marine environments.
Freshwater Biology, 48, 21–27.
Jonzén, N., Hendenström, A., and Lundberg, P. (2007a)
Climate change and the optimal arrival of migratory
birds. Proceedings of the Royal Society B, Biological
Sciences, 274, 269–74.
Jonzén, N., Ergon, T., Lindén, A., and Stenseth, N.C. (eds)
(2007b) Bird migration and climate. Climate Research
Special, 17, 1–180.
Jonzén, N., Lindén, A., Ergon, T., Knudsen, E., Vik, J.O.,
Rubolini, D., Piacentini, D., Brinch, C., Spina, F.,
Karlsson, L., Stervander, M., Andersson, A.,
Waldenström, J., Lehikoinen, A., Edvardsen, E., Solvang,
R., and Stenseth, N.C. (2006) Rapid advance of spring
arrival dates in long-distance migratory birds. Science,
312, 1959–61.
Jordano, P. (1987) Frugivory, external morphology and
digestive system in Mediterranean sylviid warblers
Sylvia spp. Ibis, 129, 175–89.
Jorgensen, S.J., Reeb, C.A., Chapple, T.K., Anderson, S.,
Perle, C., van Sommeran, S.R., Fritz-Cope, C., Brown,
A.C.; Klimley, A.P., and Block, B.A. (2010) Philopatry
and migration of Pacific white sharks. Proceedings of
the Royal Society B-Biological Sciences, 277, 679–88.
Joseph, I., Lessa, E.P., and Christidis, L. (1999) Phylogeny
and biogeography in the evolution of migration: shorebirds of the Charadrius complex. Journal of
Biogeography, 26, 329–42.
Julian, F. and Beeson, M. (1998) Estimates of marine mammal, turtle, and seabird mortality for two California
gillnet fisheries: 1990–1995. Fisheries Bulletin, 96,
271–84.
Kaartvedt, S. (1996) Habitat preferences during overwintering and timing of seasonal vertical migration of
Calanus finmarchicus. Ophelia, 44, 145–56.
Kaimal, B., Johnson, R., and Hannigan, R. (2009)
Distinguishing breeding populations of mallards (Anas
platyrhynchos) using trace elements. Journal of
Geochemical Exploration, 102, 44–48.
Kaitala, A., Kaitala, V., and Lundberg P. (1993) A theory of
partial migration. American Naturalist, 142, 59–81.
Kamp, J., Sheldon, R.D., Koshkin, M.A., Donald, P.F., and
Biedermann, R. (2009) Post-Soviet steppe management
causes pronounced synanthropy in the globally threatened Sociable Lapwing Vanellus gregarius. Ibis, 151,
452–63.
Kaplan, I.C. (2005) A risk assessment for Pacific leatherback turtles (Dermochelys coriacea). Canadian Journal of
Fisheries and Aquatic Science, 62, 1710–19.
Karasov, W.H. (1990) Digestion in birds: chemical and
physiological determinants and ecological implications.
Studies in Avian Biology, 13, 391–415.
Karasov, W.H. and Pinshow, B. (2000) Test for physiological limitation to nutrient assimilation in a long-distance
passerine migrant at a springtime stopover site.
Physiological and Biochemical Zoology, 73, 335–43.
Kareiva, P., Marvier, M., and McClure, M. (2000) Recovery
and management options for spring/summer chinook
salmon in the Columbia River Basin. Science, 290, 5493,
977–79.
Keast, A. (1968) Seasonal movements in the Australian
honeyeaters (Meliphagidae) and their ecological significance. Emu, 67, 159–209.
Keeton, W. T. (1980) Avian orientation and navigation:
new developments in an old mystery. pp. 137–58, Acta
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
XVII Congr Intern Ornithol I, Deutsche OrnithologenGesellschaft: Berlin.
Kelly, J.F., DeLay, L.S., and Finch, D.M. (2002) Densitydependent mass gain by Wilson’s Warblers during stopover. Auk, 119, 210–13.
Kennedy, J.S. (1961) A turning point in the study of insect
migration. Nature, 189, 785–91.
Kennedy, J.S. (1985) Migration, behavioural and ecological. In M.A. Rankin (ed.) Migration: Mechanisms and
Adaptive Significance, pp. 5–26, Contributions to
Marine Science, Marine Science Institute, The University
of Texas: Austin.
Kenward, R.E. (2001) A Manual for Wildlife Radiotracking,
Academic Press: London.
Kerlinger, P. and Moore, F.R. (1989) Atmospheric structure
and avian migration. Current Ornithology, 6, 109–42.
Kerven, C.K. (ed.) (2003) Prospects for Pastoralism in
Kazakhstan and Turkmenistan: From State Farms to
Private Flocks, Routledge Curzon: London.
Kerven, C.K. (2004) The influence of cold temperatures
and snowstorms on rangelands and livestock in northern Asia. In S. Vetter (ed.) Rangelands at Equilibrium
and Non-Equilibrium: Recent Developments in the
Debate around Rangeland Ecology and Management,
pp. 41–55, Programme for Land and Agrarian Studies,
University of the Western Cape: Cape Town.
Kerven, C.K., Shanbaev, K, Alimaev, I., Smailov, A., and
Smailov, K. (2008) Livestock mobility and degradation
in Kazakhstan’s semi-arid rangelands. In R. Behnke
(ed.) The Socio-Economic Causes and Consequences of
Desertification in Central Asia, pp. 113–40, Springer:
Dordrecht, The Netherlands.
Kerven, C.K., Alimaev, I.I., Behnke, R., Davidson, G.,
Malmakov, N., Smailov, A., and Wright, I. (2006)
Fragmenting pastoral mobility: changing grazing patterns in post-Soviet Kazakhstan. In D.J. Bedunah, E.D.
McArthur, and M. Fernandez-Gimenez, (comps)
Rangelands of Central Asia: Proceedings of the
Conference on Transformations, Issues, and Future
Challenges, 27 January 2004, Salt Lake City, UT.
Proceeding RMRS-P-39. U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Station: Fort
Collins, CO.
Kéry, M., Madsen, J., and Lebreton J.-D. (2006) Survival of
Svalbard pink-footed geese Anser brachyrhynchus in
relation to winter climate, density and land-use. Journal
of Animal Ecology, 75, 1172–81.
Kiffney, P.M., Pess, G.R., Anderson, J.H., Faulds, P., Burton
K., and Riley, S.C. (2009) Changes in fish communities
following recolonization of the Cedar River, WA, USA
by Pacific Salmon after 103 years of local extirpation.
River Research and Applications, 25, 438–52.
235
King, A.A. and Schaffer, W.M. (1999) The rainbow bridge:
Hamiltonian limits and resonance in predator-prey
dynamics. Journal of Mathematical Biology, 39, 439–69.
King, J.R. and Mewaldt, L.R. (1981) Variation of bodyweight in gambel white-crowned sparrows in winter
and spring—Latitudinal and photoperiodic correlates.
Auk, 98, 752–64.
Kirby, J.S., Stattersfield, A.J., Butchart, S.H.M., Evans, M.I.,
Grimmett, R.F.A., Jones, V.R., O’Sullivan, J., Tucker, G.M.,
and Newton, I. (2008) Key conservation issues for migratory land and waterbird species on the world’s major
flyways. Bird Conservation International, 18, S49–S73.
Kirilenko, A.P. and Solomon, A.M. (1998) Modeling
dynamic vegetation response to rapid climate change
using bioclimatic classification. Climatic Change, 38,
15–49.
Kitti, H., Gunslay, N., and Forbes, B.C. (2006) Defining the
Quality of Reindeer Pastures: The perspective of Sami
Reindeer Herders. In B.C. Forbes, M. Bölter, L. MüllerWille, J. Hukkinen, F. Müller, N. Gunslay, and Y.
Konstantinov (eds) Reindeer Management in Northernmost Europe, Vol. 184, Ecological Studies, pp. 141–65,
Springer: Berlin, Heidelberg.
Klaassen, M., Kvist, A., and Lindström, A. (2000) Flight
costs and fuel composition of a bird migrating in a wind
tunnel. Condor, 102, 444–51.
Klaassen, M., Bauer, S., Madsen, J., and Possingham, H.
(2008) Optimal management of a goose flyway: migrant
management at minimum cost. Journal of Applied
Ecology, 45, 1446–52.
Klaassen, M., Bauer, S, Madsen J., and Tombre, I. (2006)
Modelling behavioural and fitness consequences of disturbance for geese along their spring flyway. Journal of
Applied Ecology, 43, 92–100.
Klaassen, M., Lindström, A., Meltofte, H., and Piersma, T.
(2001) Ornithology—Arctic waders are not capital
breeders. Nature, 413, 794.
Klaassen, M., Beekman, J.H., Kontiokorpi, J., Mulder,
R.J.W., and Nolet, B.A. (2004) Migrating swans profit
from favourable changes in wind conditions at low altitude. Journal of Ornithology, 145, 142–51.
Klaassen, R.H.G. and Nolet, B.A. (2008) Persistence of spatial variance and spatial pattern in the abundance of a
submerged plant. Ecology, 89, 2973–79.
Kleist, A.M., Lancia, R.A., and Doerr, P.D. (2007) Using
Video Surveillance to Estimate Wildlife Use of a
Highway Underpass. The Journal of Wildlife Management, 71, 8, 2792–800.
Klokov, K.B. (1997) Northern reindeer of Taimyr Okrug as
the focus of economic activity: Contemporary problems
of reindeer husbandry and the wild reindeer hunt. Polar
Geography, 21, 233–71.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
236
REFERENCES
Knight, T.M., McCoy, M.W., Chase, J.M., McCoy, K.A, and
Holt, R.D. (2005) Trophic cascades across ecosystems.
Nature, 437, 880–83.
Koehler, A.V., Pearce, J.M., Flint, P.L., Franson, J.C., and Ip,
H.S. (2008) Genetic evidence of intercontinental movement of avian influenza in a migratory bird: the northern
pintail (Anas acuta) Molecular Ecology, 17, 4754–62.
Koenig, W.D. and Knops, J.M.H. (1998) Scale of mast seeding and tree-ring growth. Nature, 396, 225–26.
Kölzsch, A., and Blasius, B. (2008) Theoretical approaches
to bird migration. European Physical Journal-Special
Topics 157:191–208.
Kokko, H. (1999) Competition for early arrival in migratory birds. Journal of Animal Ecology, 68, 940–50.
Kokko, H. and Lundberg, P. (2001) Dispersal, migration,
and offspring retention in saturated habitats. The
American Naturalist, 157, 2, 188–202.
Koops, M.A. (2004) Reliability and the value of information. Animal Behaviour, 67, 103–11.
Korpimäki, E. (1986) Gradients in population fluctuations
of Tengmalm’s owl Aegolius funereus in Europe.
Oecologia, 69, 195–201.
Kram, R. and Taylor, C.R. (1990) Energetics of running: a
new perspective. Nature, 346, 265–266.
Kramer, D.L. and Chapman, M.R. (1999) Implications
of fish home range size and relocation for marine
reserve function. Environmental Biology of Fishes, 55,
65–79.
Kramer, G. (1959) Recent experiments on bird orientation.
Ibis, 101, 399–416.
Krupnik, I. (1993) Arctic Adaptations: Native Whalers and
Reindeer Herders of Northern Eurasia, University Press
of New England: Hanover, New Hampshire.
Kuha, J. (2004) AIC and BIC—Comparisons of assumptions and performance. Sociological Methods and
Research, 33, 188–229.
Kühl, A., Balinova, N., Bykova, E., Esipov, A., Arylov, I.A.,
Lushchekina, A.A., and Milner-Gulland, E.J. (2009) The
role of saiga poaching in rural communities: linkages
between attitudes, socio-economic circumstances and
behaviour. Biological Conservation, 142, 1442–1449.
Kullberg, C., Fransson, T., and Jakobsson, S. (1996)
Impaired predator evasion in fat blackcaps (Sylvia atricapilla) Proceedings of the Royal Society of London
Series B Biological Sciences, 263, 1671–75.
Kullberg, C., Henshaw, I., Jakobsson, S., Johansson, P., and
Fransson, T. (2007) Fuelling decisions in migratory
birds: geomagnetic cues override the seasonal effect.
Proceedings of the Royal Society B-Biological Sciences,
274, 2145–51.
Kumpula, T. (2006) Very High Resolution Remote Sensing
Data in Reindeer Pasture Inventory in Northern
Fennoscandia In B.C. Forbes, M. Bölter, L. Müller-Wille,
J. Hukkinen, F. Müller, N. Gunslay, and Y. Konstantinov,
Reindeer Management in Northernmost Europe, Vol.
184, Ecological Studies, pp. 167–85, Springer: Berlin,
Heidelberg.
Kunz, T.H., Wrazen, J.A., and Burnett, C.D. (1998) Changes
in body mass and fat reserves in pre-hibernating little
brown bats (Myotis lucifugus). Ecoscience, 5, 8–17.
Kvist, A. and Lindström, A. (2000) Maximum daily energy
intake: it takes time to lift the metabolic ceiling.
Physiological and Biochemical Zoology, 73, 30–36.
Kvist, A. and Lindström, A. (2003) Gluttony in migratory
waders—unprecedented energy assimilation rates in
vertebrates. Oikos, 103, 397–402.
Kvist, A., Klaassen, M., and Lindström, Å. (1998) Energy
expenditure in relation to flight speed: what is the
power of mass loss rate estimates? J. Avian Biol., 29,
485–98.
Kwan, D. (1994) Fat reserves and reproduction in the green
turtle, Chelonia mydas. Wildlife Research, 21, 257–66.
Lack, D. (1954) The Natural Regulation of Animal
Numbers, Oxford University Press: London.
Lack, D. (1968) Bird migration and natural selection.
Oikos, 19, 1–9.
Laliberte, A.S. and Ripple, W.J. (2004) Range Contractions
of North American Carnivores and Ungulates.
BioScience, 54, 2, 123–38.
Lande, R., Engen, S., and Sæther, B.-E. (2003) Stochastic
Population Dynamics in Ecology and Conservation,
Oxford Series in Ecology and Evolution, Oxford
University Press: Oxford.
Langvatn, R. and Albon, S.D. (1986) Geographic clines in
body weight of Norwegian red deer: a novel explanation of Bergmann’s rule? Holartic Ecology, 9, 285–93.
Langvatn, R. and Hanley, T.A. (1993) Feeding-patch choice
by red deer in relation to foraging efficiency. Oecologia,
95, 164–70.
Lank, D. B., Butler, R.W., Ireland, J. and Ydenberg, R. C.
(2003) Effects of predation danger on migration strategies of sandpipers. Oikos, 103, 303–320.
Lank, D.B., Pither, J., Chipley, D., Ydenberg, R.C.,
Kyser, T.K., and Norris, D.R. (2007) Trace element
profiles as unique identifiers of western sandpiper
(Calidris mauri) populations. Canadian Journal of
Zoology, 85, 579–83.
Laris, P. (2002) Burning the seasonal mosaic: preventative
burning strategies in the wooded savanna of southern
Mali. Human Ecology, 30 (2), 155–86.
Le Boeuf, B.J., Crocker, D.E., Costa, D.P., Blackwell, S.B.,
Webb, P.M., and Houser, D.S. (2000) Foraging Ecology
of Northern Elephant Seals. Ecological Monographs, 70,
353–82.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Le Houérou, H.N. (1989) The Grazing Land Ecosystems of
the African Sahel, Ecological Studies 75, SpringerVerlag: Berlin, New York.
Le Pendu, Y. and Ciofolo, I. (1999) Seasonal movements of
giraffes in Niger. Journal of Tropical Ecology, 15, 3), 341–53.
Lehikoinen, E., Sparks, T.H., and Zalakevivus, M. (2004)
Arrival and departure dates. In: A.P. Møller, W. Fiedler
and P. Berthold (eds) Birds and Climate Change.
Advances in Ecological Research 35, pp. 1–31, Elsevier:
Amsterdam, The Netherlands.
Leimgruber, P., McShea, W.J., Brookes, C.J., Bolor-Erdene,
Wemmer, L.C., and Larson, C. (2001) Spatial patterns in
relative primary productivity and gazelle migration in
the Eastern Steppes of Mongolia. Biological
Conservation, 102, 205–12.
Leon, J. (2009) Evaluating the Use of Local Knowledge in
Species Distribution Studies: A Case Study of Saiga
Antelope in Kalmykia, Russia, MSc thesis, Imperial
College: London. Available at www.iccs.org.uk.
Lepczyk, C.A., Murray, K.G., Winnett, M.K., Bartell, P., Geyer,
E., and Work, T. (2000) Seasonal fruit preferences for lipids
and sugars by American robins. Auk, 117, 709–17.
Levey, D.J. and Stiles, F.G. (1992) Evolutionary precursors
of long-distance migration: resource availability and
movement patterns in Neotropical landbirds. The
American Naturalist, 140, 447–76.
Levins, R. and Culver, D. (1971) Regional coexistence of
species and competition between rare species (mathematical model/habitable patches). Proceedings of the
National Academy of Sciences of the United States of
America, 68, 1246ff.
Lewison, R.L. and Crowder, L.B. (2007) Putting Longline
Bycatch of Sea Turtles into Perspective. Conservation
Biology, 21, 79–86.
Lewison, R.L., Freeman, S.A., and Crowder, L.B. (2004a)
Quantifying the effects of fisheries on threatened species: the impact of pelagic longlines on loggerhead and
leatherback sea turtles. Ecology, 7, 221–31.
Lewison, R.L., Crowder, L.B., Read, A.J., and Freeman,
S.A. (2004b) Understanding impacts of fisheries bycatch
on marine megafauna. Trends in Ecology and Evolution,
19, 11, 599–604.
Liechti, F. (2006) Birds: blowin’ by the wind? Journal of
Ornithology, 147, 202–11.
Liechti, F. and Bruderer, B. (1998) The relevance of wind
for optimal migration theory. Journal of Avian Biology,
29, 561–68.
Liechti, F. and Schaller, E. (1999) The use of low-level jets
by migrating birds. Naturwissenschaften, 86, 549–51.
Liechti, F., Klaassen, M., and Bruderer, B. (2000) Predicting
migratory flight altitudes by physiological migration
models. Auk, 117, 205–14.
237
Lincoln, F.C. (1939) The Migration of American Birds,
Doubleday, Doran, and Co.: New York.
Lind, J., Fransson, T., Jakobsson, S., and Kullberg, C. (1999)
Reduced take-off ability in robins (Erithacus rubecula)
due to migratory fuel load. Behavioral Ecology and
Sociobiology, 46, 65–70.
Lindström, A. (1990) The role of predation risk in stopover
habitat selection in migrating Bramblings Fringilla montifringilla. Behavioral Ecology, 1, 102–106.
Lindström, A. (1991) Maximum fat deposition rates in
migrating birds. Ornis Scandinavica, 22, 12–19.
Lindström, A. and Alerstam, T. (1992) Optimal fat loads in
migrating birds: a test of the time-minimization hypothesis. American Naturalist, 140, 477–91.
Lindström, A., Daan, S., and Visser, H.G., (1994) The conflict between moult and migratory fat deposition: a photoperiodic experiment with bluethroats. Animal
Behaviour, 48, 1173–81.
Lindström, A., Hasselquist, D., Bensch, S., and Mats, G.
(1990) Asymmetric contests over resources for survival
and migration: a field experiment with bluethroats.
Animal Behaviour, 40, 453–61.
Linnaeus, C. (1757) Migrationes Avium.
Löft, E.R., Menke, J.W., and Burton, T.S. (1984) Seasonal
movements and summer habits of female black-tailed
deer. Journal of Wildlife Management, 48, 1317–25.
Lohmann, K.J. and Lohmann, C.M.F. (1996) Orientation
and open-sea navigation in sea turtles. Journal of
Experimental Biology, 199, 73–81.
Lohmann, K.J., Hester, J.T. and Lohmann, C.M.F. (1999)
Long-distance navigation in sea turtles. Ethology,
Ecology and Evolution, 11, 1–23.
Lohmann, K.J., Lohmann, C.M.F., and Endres, C.S. (2008a)
The sensory ecology of ocean navigation. Journal of
Experimental Biology, 211, 1719–28.
Lohmann, K.J., Luschi, P., and Hays, G.C. (2008b) Goal
navigation and island-finding in sea turtles. Journal of
Experimental Marine Biology and Ecology, 356,
83–95.
Lohmann, K.J., Putman, N.F., and Lohmann, C.M.F.
(2008) Geomagnetic imprinting: a unifying hypothesis of long-distance natal homing in salmon and sea
turtles. Proceedings of the National Academy of
Sciences of the United States of America, 105, 49,
19096–101.
Lohmann, K.J., Cain, S.D., Dodge, S.A., and Lohmann,
C.M.F. (2001) Regional magnetic fields as navigational
markers for sea turtles. Science, 294, 364–66.
Loonen, M., Zijlstra, M., and Vaneerden, M.R. (1991)
Timing of wing molt in greylag geese Anser anser in relation to the availability of their food plants. Ardea,
79:252–59.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
238
REFERENCES
Loreau, M., Mouquet, N.M., and Holt, R.D. (2005) From
metacommunities to metaecosystems. In M. Holyoak,
M.A. Leibold, and R.D. Holt (eds) Metacommunities:
Spatial Dynamics and Ecological Communities,
pp. 418–38, University of Chicago Press: Chicago.
Louchart, A. (2008) Emergence of long distance bird
migrations: a new model integrating global climate
changes. Naturwissenschaften, 95, 1109–19.
Lovejoy, N.R., Mullen, S.P., Sword, G.A., Chapman, R.F.,
and Harrison, R.G. (2006) Ancient trans-Atlantic flight
explains locust biogeography: molecular phylogenetics
of Schistocerca. Proceedings of the Royal Society B,
Biological Sciences, 273, 767–74.
Lumsden, H.G. (1984) The pre-settlement breeding distribution of Trumpeter (Cygnus buccinator) and Tundra
swans (Cygnus columbianus) in Eastern Canada.
Canadian Field-Naturalist, 98, 415–24.
Lumsden, H.G. and Drever, M.C. (2002) Overview of the
Trumpeter Swan Reintroduction Program in Ontario,
1982–2000. Waterbirds: The International Journal of
Waterbird Biology, 25, Special Publication 1, Proceedings
of the Fourth International Swan Symposium, 2001,
pp. 301–12.
Lundberg, J. and Moberg, F. (2003) Mobile link organisms
and ecosystem functioning: Implications for ecosystem
resilience and management. Ecosystems, 6, 87–98.
Lundberg, P. (1987) Partial bird migration and evolutionary stable strategies. Journal of Theoretical Biology, 125,
351–60.
Lundberg, P. (1988) The evolution of partial migration in
birds. Trends in Ecology and Evolution, 3, 172–75.
Luschi, P., Hays, G.C., and Papi, F. (2003) A review of longdistance movements by marine turtles, and the possible
role of ocean currents. Oikos, 103, 293–302.
Luschi, P., Hays, G.C., del Seppia, C., Marsh, R., and Papi,
F. (1998) The navigational feats of green sea turtles
migrating from Ascension Island investigated by satellite telemetry. Proceedings of the Royal Society of
London Series B-Biological Sciences, 265, 2279–84.
Luschi, P., Åkesson, S., Broderick, A.C., Glen, F., Godley,
B.J., Papi, F. and Hays, G.C. (2001) Testing the navigational abilities of ocean migrants: displacement experiments on green sea turtles (Chelonia mydas) Behavioral
Ecology and Sociobiology, 50, 528–34.
Lushchekina, A.A. and Struchkov, A. (2001) The saiga
antelope in Europe: Once again at the brink? The Open
Country, 3, 11–24.
Lyons, J.E., Collazo, J.A., and Guglielmo, C.G. (2008)
Plasma metabolites and migration physiology of semipalmated sandpipers: refueling performance at five latitudes. Oecologia, 155, 417–27.
Mace, G.M. and Reynolds, J.D. (2001) Exploitation as a
conservation issue. In J.D. Reynolds, G.M. Mace, K.H.
Redford, and J.G. Robinson (eds) Conservation of
Exploited Species, pp. 3–15, Cambridge University
Press: Cambridge.
Maddock, L. (1979) The ‘migration’ and grazing succession. In A.R.E. Sinclair and M. Norton-Griffiths (eds)
Serengeti: Dynamics of an Ecosystem, pp. 104–29,
University of Chicago Press: Chicago.
Magnus, O. (1555) Historia de Gentibus Septenionalibus.
Mahan, T.A. and Simmers, B.S. (1992) Social preference of
four cross-foster reared Sandhill Cranes. Proceedings of
the North American Crane Workshop, 6, 114–119.
Mahoney, S.P. and Schaefer, J.A. (2002) Hydroelectric
development and the disruption of migration in caribou. Biological Conservation, 107, 147–53.
Mahoney, S.P. and Schaefer, J.A. (2002) Long-term changes
in demography and migration of Newfoundland caribou. Journal of Mammalogy, 83, 957–63.
Maj, E. (2009) La vache sedentaire, le renne et le cheval
nomades chez les Evenes et las Iakoutes des monts de
Verkhihansk (Republique Sakha, Iakoutie). Fondation
Fyssen—Annales, 23, 36–48.
Malcolm, J.R., Markham, A., Neilson, R.P., and Garaci, M.
(2002) Estimated migration rates under scenarios of global climate change. Journal of Biogeography, 29,
835–49.
Malechek, J.C. (1984) Impacts of grazing intensity and specialized grazing systems on livestock response. In
Developing Strategies for Rangeland Management, pp.
1129–58, National Academy of Sciences and Westview
Press: Boulder, Colorado and London.
Mallon, D.P. and Kingswood, S.C. (compilers) (2001)
Antelopes, Part 4: North Africa, the Middle East, and
Asia, Global Survey and Regional Action Plans, SSC
Antelope Specialist Group, IUCN:Gland, Switzerland
and Cambridge, UK.
Mallory, F.F. and Hillis, T.L. (1998) Demographic characteristics of circumpolar caribou populations: ecotypes,
ecological constraints, releases, and population dynamics. Rangifer, Special Issue, 10, 49–60.
Mandel, J.T., Bildstein, K.L., Bohrer, G., and Winkler, D.W.
(2008) The movement ecology of migration in Turkey
Vultures. Proceedings of the National Academy of Sciences
of the United States of America, 105, 19102–19107.
Mangel, M. (1994) Climate change and salmonid life history variation. Deep Sea Research II, 41, 75–106.
Mangel, M. and Clark, C.W. (1988) Dynamic Modeling in
Behavioral Ecology, Princeton University Press:
Princeton, New Jersey.
Mangel. M. and Satterthwaite, W.H. (2008) Combining
proximate and ultimate approaches to understand life
history variation in salmonids with application to fisheries, conservation, and aquaculture. Bulletin of Marine
Science, 83, 107–30.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Marcovaldi, M.Â., Baptistotte, C., De Castilhos, J.C., Gallo,
B.M.G., Lima, E.H.S.M., Sanches, T.M., and Vieitas, C.F.
(1998) Activities by Project TAMAR in Brazilian sea turtle feeding grounds. Marine Turtle Newsletter, 80L,
5–7.
Marks, J.C., Haden, G.A., O’Neill, M and Pace, C. (in press)
Effects of flow restoration and exotic species removal on
recovery of native fish: lessons from a dam decommissioning. Restoration Ecology, http://dx.doi.org/10.1111/
j.1526-100X.2009.00574.x
Marra, P.P., Francis, C.M., Mulvihill, R.S., and Moore, F.R.
(2005) The influence of climate on the timing and rate of
spring bird migration. Oecologia, 142, 307–15.
Marsh, R.E., Erickson, W.A., and Salmon, T.P. (1992)
Scarecrows and Predator Models for Frightening Birds
from Specific Areas. In J. E. Borrecco and R. E. Marsh
(eds) Proceedings of the Fifteenth Vertebrate Pest
Conference, University of Nebraska, Lincoln, University
of California: Davis.
Marshall, F. (1990) Origins of specialized pastoral production in East Africa. American Anthropologist, 92, 873–94.
Martin, T.G., Chadès, I., Arcese, P., Marra, P.P., Possingham,
H.P., and Norris, D.R. (2007) Optimal Conservation of
Migratory Species. Public Library of Science One, 2, E751.
Marty, A. (1993) La gestion des terroirs et les éleveurs: Un
outil d’exclusion ou de négociation? Revue Tiers Monde,
34, 134, 329–44.
Matthiopoulos, J., Harwood, J., and Thomas, L. (2005)
Metapopulation consequences of site fidelity for colonially breeding mammals and birds. Journal of Animal
Ecology, 74, 716–27.
Matthiopoulos, J., Thomas, L., McConnell, B., Duck, C.,
Thompson D., Pomeroy, P., Harwood, J., MilnerGulland, E.J., Wolf, N., and Mangel, M. (2008) Putting
Long-Term, Population Monitoring Data to Good Use:
The Causes of Density Dependence in UK grey seals.
SCOS Briefing paper 06/08.
May, R.M. (1976) Simple models with very complicated
dynamics. Nature, 261, 459–67.
Maynard-Smith, J. (1982) Evolution and the Theory of
Games, Cambridge University Press: Cambridge.
Mayr, E. (1961) Cause and Effect in Biology. Science, 134,
1501–1506.
Mayr, E. and Meise, W. (1930) Theoretisches zur Geschichte
des Vogulzuges. Der Vogelzug (Berlin), 1, 149–72.
Mbaiwa, J. E. and Darkoh, M.B.K (2005) Sustainable development and natural resource competition and conflicts
in the Okavango Delta, Botswana. Botswana Notes and
Records, 37, 40–60.
Mbaiwa, J.E. and Mbaiwa, O.I. (2006) The effects of veterinary fences on wildlife populations in Okavango Delta,
Botswana. International Journal of Wilderness, 12, 3,
17–23.
239
McCabe, J.T. (1983) Land use among the pastoral Turkana.
Rural Africana, 15–16, 109–26.
McCabe, J.T. (1994) Mobility and land use among African
pastoralists: old conceptual problems and new interpretations. In E. Fratkin, K.A. Galvin, and E.A. Roth (eds)
African Pastoralist Systems: An Integrated Approach,
pp. 69–89, Lynne Rienner Publishers: Boulder.
McClelland, G.B. (2004) Fat to the fire: the regulation of
lipid oxidation with exercise and environmental stress.
Comparative Biochemistry and Physiology, Part B, 139,
443–60.
McClenachan, L., Jackson, J.B.C., and Newman, M.J.H.
(2006) Conservation Implications of Historic Sea Turtle
Nesting Beach Loss. Frontiers in Ecology and the
Environment, 4, 6, 290–96.
McConville, A.J., Grachev, I.A., Keane, A., Coulson, T.,
Bekenov, A., and Milner-Gulland, E.J. (2009) Reconstructing the observation process to correct for changing
detection probability of a critically endangered species.
Endangered Species Research, 6, 231–37.
McCormick, S.D., Hansen, L.P., Quinn, T.P., and Saunders,
R.L. (1998) Movement, migration, and smolting of
Atlantic salmon (Salmo salar). Canadian Journal of
Fisheries and Aquatic Sciences, 55, Suppl. 1, 77–92.
McEneaney, T. (2001) Yellowstone Bird Report, 2000.
National Park Service, Yellowstone Center for Resources,
Yellowstone National Park: Wyoming, YCR–NR–2001–01.
McGuire, L.P. and Guglielmo, C.G. (2009) What can birds
tell us about the migration physiology of bats? Journal
of Mammalogy, 90, 1290–97.
McKelvey, K.S. and Noon, B.R. (2001) Incorporating uncertainties in animal location and map classification into
habitat relationships modeling. In C. T. Hunsaker,
M. F. Goodchild, M. A. Friedl, and T. J. Case (eds) Spatial
Uncertainty in Ecology, pp. 72–90, Springer-Verlag:
New-York.
McKenzie, J. (2001) The selective advantage of urban habitat use by elk in Banff National Park M.Sc. Thesis,
University of Guelph: Ontario.
McKinnon, L., Smith, P.A., Nol, E., Martin, J.L., Doyle, F.I.,
Abraham, K.F., Gilchrist, H.G., Morrison, R.I.G., and
Bety, J. (2010) Lower predation risk for migratory birds
at high latitudes. Science, 327, 326–27.
McLellan, B.N. and Shackleton, D.M. (1988) Grizzly bears
and resource-extraction industries: effects of roads on
behavior, habitat use, and demography. Journal of
Applied Ecology, 25, 451–60.
McNamara, J.M. and Houston, A.I. (2008) Optimal annual
routines: behaviour in the context of physiology and
ecology. Philosophical Transactions of the Royal Society
B, 363, 301–19.
McNaughton, S.J. (1976) Serengeti migratory wildebeest:
facilitation of energy flow by grazing. Science, 191, 92–4.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
240
REFERENCES
McNaughton, S.J. (1984) Grazing lawns: animals
in herds, plant form, and coevolution. American
Naturalist, 124, 863–86.
McPeek, M.A. and Holt, R.D. (1992) The evolution
of dispersal in spatially and temporally varying
environments. American Naturalist, 140, 1010–27.
McQuire, L.P. and Guglielmo, C.G. (2009) What can birds
tell us about the migration physiology of bats? Journal
of Mammalogy, 90, 1290–97.
McWilliams, S.R., Kearney, S.B., and Karasov, W.H. (2002)
Diet preferences of warblers for specific fatty acids in
relation to nutritional requirements and digestive capabilities. Journal of Avian Biology, 33, 167–74.
Mduma, S.A.R., Sinclair, A.R.E., and Hilborn, R. (1999)
Food regulates the Serengeti wildebeest: a 40-year
record. Journal of Animal Ecology, 68, 1101–22.
Mehlman, D.W., Mabey, S.E., Ewert, D.N., Duncan, C.,
Abel, B., Cimprich, D., Sutter, R., and Woodfrey, M.
(2005) Conserving stop-over sites for forest-dwelling
migratory landbirds. Auk, 122, 1–11.
Meitner, C.J., Brower, L.P., and Davis, A.K. (2004) Migration
patterns and environmental effects on stopover of
monarch butterflies (Lepidoptera, Nymphalidae) at
Peninsular Point, Michigan. Environmental Entomology,
33, 249–56.
Menu, S., Gauthier, G., and Reed, A. (2005) Survival of
young greater snow geese (Chen caerulescens atlantica)
during fall migration. The Auk 122, 479–96.
Mikkola, K. (2003) Red Admirals Vanessa atalanta
(Lepidoptera: Nymphalidae) select northern winds on
southward migration. Entomol Fenn, 14, 15–24.
Milá, B., Smith, B.S., and Wayne, R.K. (2006) Postglacial
population expansion drives the evolution of long-distance migration in a songbird. Evolution, 60, 2403–409.
Mills, M.G.L. and Funston, P. (2003) Large carnivores and
savannah heterogeneity In J. Du Toit, K.H. Rogers, and
H.C. Biggs (eds) The Kruger Experience, pp. 370–88,
Island Press: Washington.
Millspaugh, J.J. and Marzluff, J.M. (2001), Radio Tracking
and Animal Populations, Academic Press: San Diego.
Milner-Gulland, E.J. (2001) A dynamic game model for the
decision to join an aggregation. Ecological Modeling,
145, 85–99.
Milner-Gulland, E.J., Bukreeva, O.M., Coulson, T.N.,
Lushchekina, A.A., Kholodova, M.V., Bekenov, A.B.,
and Grachev, Iu.A. (2003) Reproductive collapse in saiga
antelope harems. Nature, 422, 135.
Milner-Gulland, E.J., Kholodova, M.V., Bekenov, A.,
Bukreeva, O.M., Grachev, Iu.A., Amgalan, L., and
Lushchekina, A.A. (2001) Dramatic declines in Saiga
antelope populations. Oryx, 35, 340345.
Minetti, A.E. (1995) Optimum gradient of mountain paths.
Journal of Applied Physiology, 79, 1698–1703.
Mitchell, C.D. (1994) Trumpeter swan (Cygnus buccinator).
In A. Poole (ed.) The Birds of North America, Online,
[Online], Available: http://bna.birds.cornell.edu.subzero.lib.uoguelph.ca/bna/species/105 [December 2008],
Cornell Lab of Ornithology, Ithaca.
Mitchell-Olds, T., Feder, M., and Wray, G. (2008)
Evolutionary and ecological functional genomics.
Heredity, 100, 101–102.
Mitcheson, Y.S.D., Cornish, A, Domeier, M, Colin, P.L.,
Russell, M., and Lindeman, K.C. (2008) A Global
Baseline for Spawning Aggregations of Reef Fishes.
Conservation Biology, 22, 5, 1233–44.
Moil, R.J., Millspaugh, J.J. Beringer, J. Sartwell, J., and He,
Z.H. (2007) A new ‘view’ of ecology and conservation
through animal-borne video systems. Trends in Ecology
and Evolution, 22, 660–68.
Møller, A.P., Rubolini, D., and Lehikoinen, E. (2008)
Populations of migratory bird species that did not show
a phenological response to climate change are declining.
Proceedings of the National Academy of Sciences of the
United States of America, 105, 42, 16195–200.
Mooney-Seus, M.L. and Rosenberg, A.A. (2007) Regional
Fisheries Management Organizations (RFMOs):
Progress in Adopting Precautionary Approach and
Ecosystem-Based Management [Online], Available:
at:http://www.illegal-fishing.info/uploads/RFMOreport-FortHill-0207.pdf [January 2009].
Moorcroft, P.R. and Lewis, M.A. (2006), Mechanistic Home
Range Analysis: Monographs in Population Biology,
Princeton University Press: Princeton.
Moore, A., Freake, S.M., and Thomas, I.M. (1990) Magnetic
particles in the lateral line of the Atlantic salmon (Salmo
salar L). Philosophical Transactions of the Royal Society
of London, 329, 11–15.
Moore, F.R. and Yong, W. (1991) Evidence of food-based
competition among passerine migrants during stopover. Behavioral Ecology and Sociobiology, 28,
85–90.
Morales, J.M., D.T. Haydon, J. Frair, K.E. Holsiner, and
J.M. Fryxell. (2004) Extracting more out of relocation
data: Building movement models as mixtures of random walks. Ecology, 85, 2436–45.
Moreau, R.E. (1972) The Palaearctic—African Bird Migration Systems, Academic Press: London, New York.
Morgan, E.R., Lundervold, M., Medley, G.F., Shaikenov,
B.S., Torgerson, P., and Milner-Gulland, E.J. (2006)
Assessing risks of disease transmission between wildlife and livestock: The Saiga antelope as a case study.
Biological Conservation, 131, 244–54.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Morris, S. R. (1996) Mass loss and probability of stopover
by migrant warblers during spring and fall migration.
Journal of Field Ornithology, 67, 456–62.
Morrison, R.I.G., Davidson, N.C., and Wilson, J.R. (2007)
Survival of the fattest: body stores on migration and
survival in red knots Calidris canutus islandica. Journal of
Avian Biology, 38, 4, 479–87.
Mortimer, J. A. and Carr, A. (1987) Reproduction and
migrations of the Ascension-Island green turtle
(Chelonia-Mydas). Copeia 1987, 103–13.
Mosser, A., and Packer, C. (2009) Group territoriality and
the benefits of sociality in the African lion, Panthera leo,
Animal Behaviour 78, 359–70.
Mouritsen, H. and Frost, B. J. (2002) Virtual migration in
tethered flying monarch butterflies reveals their orientation mechanisms. Proceedings of the National Academy
of Sciences USA, 99, 10162–66.
Mueller, T. and Fagan, W. F. (2008) Search and navigation
in dynamic environments—from individual behaviors
to population distributions. Oikos, 117, 654–64.
Mueller, T., Olson, K.A., Fuller, T.K., Schaller, G.B., Murray
M.G., and Leimgruber, P. (2008) In search of forage: predicting dynamic habitats of Mongolian gazelles using
satellite-based estimates of vegetation productivity.
Journal of Applied Ecology, 45, 649–58.
Muheim, R., Phillips, J.B. and Åkesson, S. (2006) Polarized
light cues underlie compass calibration in migratory
songbirds. Science, 313, 837–39.
Murray, M.G. and Brown, D. (1993) Niche separation of
grazing ungulates in the Serengeti—an experimental
test. Journal of Animal Ecology, 62, 380–89.
Murray, M.G. and Illius, AW. (1996) Multispecies grazing
in the Serengeti. In J. Hodgson and A.W. Illius (eds)
The Ecology and Management of Grazing Systems,
pp. 247–74, CAB International: Wallingford, UK.
Murray, M.G. and Illius, AW. (2000) Vegetation modification and resource competition in grazing ungulates.
Oikos, 89, 501–508.
Murtaugh, P.A. (2009) Performance of several variable-selection methods applied to real ecological data. Ecology
Letters, 12, 1061–68.
Myers, G.S. (1949) Usage of anadromous, catadromous
and allied terms for migratory fishes. Copeia, 1949,
89–97.
Nakazawa, Y., Peterson, A.T., Martinez-Meyer, E., and
Navarro-Siguenza, A.G. (2004) Seasonal niches of
Nearctic–Neotropical migratory birds: Implications for
the evolution of migration. Auk, 121, 610–18.
Nathan, R., Getz, W.M., Revilla, E., Holyoak, M., Kadmon,
R., Saltz, D., and Smouse, P.E. (2008) A movement ecology paradigm for unifying organismal movement
241
research. Proceedings of the National Academy of
Sciences USA, 105, 19052–59.
Nellemann, C., Vistnes, I., Jordhøy, P., Støen, O.-G.,
Kaltenborn, B.P., F. Hanssen, and Helgesen, R. (2009)
Effects of recreational cabins, trails and their removal
for restoration of reindeer winter ranges. Restoration
Ecology, 17, 1, 1–9.
Nelson, M.E. (1998) Development of migratory behavior
in northern white-tailed deer. Canadian Journal of
Zoology, 76, 426–32.
Nelson, M.E. and Mech, L.D. (1981) Deer Social
Organization and Wolf Predation in Northeastern
Minnesota, Wildlife Monographs, No.77, 1–53.
Newton, I. (2006) Advances in the study of irruptive
migration. Ardea, 94, 433–60.
Newton, I. (2008) The Migration Ecology of Birds,
Academic Press: Oxford.
Newton, I. and Dale, L.C. (1996a) Bird migration at different
latitudes in eastern North America. Auk, 113, 626–35.
Newton, I. and Dale, L.C. (1996b) Relationship between
migration and latitude among west European birds.
Journal of Animal Ecology, 65, 137–46.
Ng, S.J., Dole, J.W., Sauvajot, R.M., Riley, S.P.D., and
Valonec, T.J. (2004) Use of highway undercrossings by
wildlife in southern California. Biological Conservation,
115, 499–507.
Niamir-Fuller, F. (1998) The resilience of pastoral herding
in Sahelian Africa. In F. Berkes, C. Folke, and J. Colding
(eds) Linking social and ecological systems: management practices and social mechanisms for building
resilience, pp. 250–84, Cambridge University Press:
Cambridge, UK.
Nilsen, T.O., Ebbesson, L.O.E., Madsen, S.S., McCormick,
S.D. Andersson, E., BjÖrnsson, B.T., Prunet, P., and
Stefansson, S.O. (2007) Differential expression of gill
Na+,K+- ATPase alpha- and beta-subunits, Na+,K+,2Cl(-)
cotransporter and CFTR anion channel in juvenile
anadromous and landlocked Atlantic salmon Salmo
salar. Journal of Experimental Biology, 210, 2885–96.
Nilsson, A.L.K., Lindstrom, A., Jonzén, N., Nilsson, S.G.,
and Karlsson, L. (2006) The effect of climate change on
partial migration—the blue tit paradox. Global Change
Biology, 12, 10, 2014–22.
Nisbet, I.C.T. (1955) Atmospheric turbulence and bird
flight. British Birds, 48, 557–59.
Nisbet, I.C.T. and Drury, W.H. (1967) Orientation of spring
migrants studied by radar. Bird Banding, 38, 173–86.
Nocera, J. J., Taylor, P.D., and Ratcliffe, L.M. (2008) Inspection
of mob-calls as sources of predator information: response
of migrant and resident birds in the Neotropics.
Behavioural Ecology and Sociobiology, 62, 1769–77.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
242
REFERENCES
Nolet, B.A. (2006) Speed of spring migration of Tundra
Swans Cygnus columbianus in accordance with income
or capital breeding strategy? Ardea, 94, 579–91.
Norling, B.S. (1992) Roost sites used by Sandhill Crane
staging along the Platte River, Nebraska. Great Basin
Naturalist, 52, 3, 253–61.
Norris, D.R. and Taylor, C.M. (2006) Predicting the consequences of carry-over effects for migratory populations.
Biology Letters, 2, 148–51.
Norris, D.R., Marra, P.P., Kyser, T.K., and Ratcliffe, L.M.
(2005) Tracking habitat use of a long-distance migratory
bird, the American redstart Setophaga ruticilla, using
stable-carbon isotopes in cellular blood. Journal of Avian
Biology, 36, 164–70.
Norton-Griffiths, M. (2007) How many wildebeest do you
need? World Economics, 8, 2, 41–64.
Norton-Griffiths, M. and Southey, C. (1995) The opportunity costs of biodiversity conservation in Kenya.
Ecological Economics, 12, 125–39.
Nøttestad, L., Giske, J., Holst, J.C. and Huse, G. (1999)
A length-based hypothesis for feeding migrations
in pelagic fish. Canadian Journal of Fisheries and
Aquatic Sciences, 56, 26–34.
Nowakowski, J.K., Remisiewicz, M., Keller, M., Busse P.,
and Rowiński, P. (2005) Synchronisation of the autumn
mass migration of passerines: a case of Robins Erithacus
rubecula. Acta Ornithologica, 40, 103–15.
Ohashi, K., Leslie, A., and Thomson, J.D. (2008) Trapline
foraging by bumble bees: V. Effects of experience and
priority on competitive performance. Behavioral
Ecology, 19, 5, 936–948.
Ohashi, K. and Thomson, J.D. (2005) Efficient harvesting
of renewing resources. Behavioral Ecology, 16, 592–605.
Okayasu, T., Muto, M., Jamsran, U., and Takeuchi K. (2007)
Spatially heterogeneous impacts on rangeland after
social system change in Mongolia. Land Degradation
and Development, 18, 5, 555–66.
Økland, F., Jonsson, B., Jensen, A.J., and Hansen, L.P.
(1993) Is there a threshold size regulating seaward
migration of brown trout and Atlantic salmon? Journal
of Fish Biology, 42, 541–50.
Okoti, M., Ng’ethe, J.C., Ekaya, W.N., and Mbuvi, D.M.
(2004) Land use, ecology, and socio-economic changes
in a pastoral production system. Journal of Human
Ecology, 16, 2, 83–9.
Oliveira, E.G., Srygley, R.B. and Dudley, R. (1998) Do neotropical migrant butterflies navigate using a solar compass? Journal of Experimental Biology, 201, 3317–31.
Olsen, J.B., Wuttig, K., Fleming, D., Kretschmer, E.J., and
Wenburg, J.K. (2006) Evidence of partial anadromy and
resident-form dispersal bias on a fine scale in populations of Oncorynchus mykiss. Conservation Genetics, 7,
613–19.
Osborn, F.V. and Parker, G.E. (2003) Linking two elephant refuges with a corridor in the communal lands
of Zimbabwe. African Journal of Ecology, 41, 1,
68–74.
Ouellet, J.-P., Crëte, M., Maltais, J, Pelletier, C., and Huot,
J. (2001) Emergency feeding of white-tailed deer: test of
three feeds. Journal of Wildlife Management, 65,
129–36.
Outlaw, D.C. and Voelker, G. (2006) Phylogenetic tests of
hypotheses for the evolution of avian migration: a case
study using the Motacillidae. The Auk, 123, 455–66.
Owen, M. and Black, J.M. (1991) A note on migration mortality and its significance in goose populations dynamics. Ardea, 79, 195–96.
Owen, M. and Gullestad, N. (1984). Migration routes of
Svalbard barnacle geese, Branta leucopsis, with a preliminary report on the importance of the Bjørnøya staging
area. Norsk Polarinstitutt Skrifter, 81, 67–77.
Owen-Smith, N. and P. Novellie (1982) What should a
clever ungulate eat? The American Naturalist, 119, 2,
151–178.
Owen-Smith, N. and Ogutu, J. (2003) Rainfall influences
on ungulate population dynamics. In J. Du Toit, K.H.
Rogers and H.C. Biggs (eds) The Kruger Experience, pp.
310–31, Island Press: Washington.
Packer, C., Hilborn, R., Mosser, A., Kissui, B., Borner, M.,
Hopcraft, G., Wilmshurst, J., Mduma, S, and Sinclair,
ARE. (2005) Ecological change, group territoriality, and
population dynamics in Serengeti lions. Science, 307,
390–93.
Palacín, C., Alonso, J.C., Alonso, J.A., Martín, C.A.,
Magaña, M., and Martin, B. (2009) Differential migration by sex in the Great Bustard: possible consequences
of an extreme sexual size dimorphism. Ethology, 115,
617–26.
Papi, F., Luschi, P., Åkesson, S., Capogrossi, S., and Hays,
G.C. (2000) Open-sea migration of magnetically disturbed sea turtles. Journal of Experimental Biology, 203,
3435–43.
Parmesan C. (2003) Butterflies as bio-indicators of climate
change impacts. In C.L. Boggs, W.B. Watt and P.R.
Ehrlich (eds) Evolution and Ecology Taking Flight:
Butterflies as Model Systems, pp. 541–60, University of
Chicago Press: Chicago.
Parmesan, C. and Yohe, G. (2003) A globally coherent fingerprint of climate change impacts across natural
systems. Nature, 421, 37–42.
Pärt, T. (1995) The importance of local familiarity and
search costs for age- and sex-biased philopatry in the
collared flycatcher. Animal Behaviour, 49, 4,
1029–38.
Parvinen, K. (1999). Evolution of migration in a metapopulation. Bulletin of Mathematical Biology, 61, 531–50.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Pascual, M.A., Bentzen, P., Riva Rossi, C., Mackey, G.,
Kinnison, M., and Walker, R. (2001) First documented
case of anadromy in a population of introduced rainbow trout in Patagonia, Argentina. Transactions of the
American Fisheries Society, 130, 53–67.
Patterson, T.A., Thomas, L., Wilcox, C., Ovaskainen O.,
and Matthiopoulos, J. (2008) State-space models of individual animal movement. Trends in Ecology and
Evolution, 23, 87–94.
Pavey, C.R. and Nano, C.E.M. (2009) Bird assemblages of
arid Australia: vegetation patterns have a greater effect
than disturbance and resource pulses. Journal of Arid
Environments, 73, 634–42.
Pegg, M.A., Layzer, J.B., and Bettoli, P.W. (1996) Angler
exploitation of anchor-tagged saugers in the lower
Tennessee River. North American Journal of Fisheries
Management, 16, 218–22.
Pelto, P. J. (1987) The Snowmobile Revolution. Technology
and Social Change in the Arctic. Prospect Heights,
Waveland Press: Illinois.
Pener M.P. and Simpson S.J. (2009) Locust phase polyphenism: An update. Advances in Insect Physiology, 36,
1–272.
Penning de Vries, F.W.T. and Djitèye, MA. (eds) (1982) La
productivité des pâturages sahéliens, Centre for
Agricultural Publishing and Documentation: Wageningen,
The Netherlands.
Pennycuick, C. J. (1969) The mechanics of bird migration.
Ibis, 111, 525–56.
Pennycuick, C. J. (1972) Animal Flight, Edward Arnold:
London.
Pennycuick, C.J. (1978) Fifteen testable predictions about
bird flight. Oikos, 30, 165–76.
Pennycuick, C.J. (1997) Actual and ‘optimum’ flight
speeds: Field data reassessed. Journal of Experimental
Biology, 200, 2355–61.
Pennycuick, C. J. (1998) Computer simulation of fat and
muscle burn in long-distance bird migration. Journal of
Theoretical Biology, 191, 47–61.
Pennycuick, C.J. (1998) Field observations of thermals and
thermal streets, and the theory of cross-country soaring
flight. Journal of Avian Biology, 29, 33–43.
Pennycuick, C.J., Einarsson, O., Bradbury, T.A.M., and
Owen, M. (1996) Migrating Whooper Swans Cygnus
cygnus: satellite tracks and flight performance calculations. Journal of Avian Biology, 27, 118–34.
Pennycuick, L. (1975) Movements of the migratory wildebeest population in the Serengeti areas between 1960
and 1973. East African Wildlife Journal, 13, 65–87.
Percival, D.P. and Walden, A.T. (2000) Wavelet methods
for Time Series Analysis, Cambridge Series in Statistical
and Probabilistic Mathematics, Cambridge University
Press: Cambridge, UK.
243
Perdeck, A.C. (1967) Orientation of starlings after displacement to Spain. Ardea, 55, 194.
Peres, C.A. (2005) Why we need megareserves in
Amazonia. Conservation Biology, 19, 3, 728–33.
Pess, G.R., McHenry, M.L, Beechie, T.J., and Davies, J.
(2008) Biological impacts of the Elwha River dams and
potential salmonid responses to dam removal. Northwest Science, 82, Special Issue, 72–90.
Peterson, C. and Messmer, T.A. (2006) Effects of winterfeeding on mule deer in Northern Utah. The Journal of
Wildlife Management, 71, 5, 1440–45.
Petersons, G. (2004) Seasonal migrations of north-eastern
populations of Nathusius’ bat Pipistrellus nathusii
(Chiroptera). Myotis, 41–42, 29–56.
Petrie, S.A. and Wilcox, K.L. (2003) Migration chronology
of Eastern-population tundra swans. Canadian Journal
of Zoology/Revue Canadien de Zoologie 81, 861–70.
Pettifor, R.A., Caldow, R.W.G., Rowcliffe, J.M., GossCustard, J.D., Black, J.M., Hodder, K.H., Houston, A.I.,
Lang, A. and Webb, J. (2000) Spatially explicit, individual-based, behavioural models of the annual cycle of
two migratory goose populations. Journal of Applied
Ecology, 37, 103–35.
Pettorelli, N., Vik, J.O., Mysterud, A., Gaillard, J.-M.,
Tucker, C.J. and Stenseth, N.C. (2005) Using the satellitederived NDVI to assess ecological responses to environmental change. Trends in Ecology and Evolution, 20,
503–10.
Philips, A.R. (1975) The migration of Allen’s and other
hummingbirds. Condor, 77, 196–205.
Piersma, T. (1998) Phenotypic flexibility during migration:
optimization of organ size contingent on the risks and
rewards of fueling and flight? Journal of Avian Biology,
29, 511–20.
Piersma, T. and Gill, R.E. (1998) Guts don’t fly: Small
digestive organs in obese Bar-tailed Godwits. Auk, 115,
196–203.
Piersma, T., Koolhaas, A., and J. Jukema (2003) Seasonal
body mass changes in Eurasian golden plovers Pluvialis
apricaria staging in the Netherlands: decline in late
autumn mass peak correlates with increase in raptor
numbers. Ibis, 145, 565–71.
Piersma, T., Pérez-Tris, J., Mouritsen, H., Bauchinger, U.,
and Bairlein, F. (2005) Is there a ‘migratory syndrome’
common to all migrant birds? Annals of the New York
Academy of Sciences, 1046, 282–93.
Pigliucci, M. (2003) Phenotypic integration: studying the
ecology and evolution of complex phenotypes. Ecology
Letters, 6, 265–72.
Pigliucci, M. and Murren, C.J. (2003) Genetic assimilation
and a possible evolutionary paradox: can macroevolution sometimes be so fast as to pass us by? Evolution, 57,
1455–64.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
244
REFERENCES
Pimm, S.L. and Raven, P. (2000) Extinction by numbers.
Nature, 403, 843-845.
Pinto, N. and Keitt, T.H. (2009) Beyond the least-cost
path: evaluating corridor redundancy using a
graph-theoretic approach. Landscape Ecology, 24,
253–66.
Poche, R.M. (1974) Notes on the roan antelope (Hippotragus
equinus (Desmarest)) in West Africa. Journal of Applied
Ecology, 11, 3, 963–68.
Poiner, I.R. and Harris, A.N.M. (1996) Incidental capture,
direct mortality and delayed mortality of sea turtles in
Australia’s northern prawn fishery. Marine Biology, 125,
813–25.
Polis, G.A., Anderson, W.B., and Holt, R.D. (1997) Toward
an integration of landscape and food web ecology: the
dynamics of spatially subsidized food webs. Annual
Review of Ecology and Systematics, 28, 289–316.
Polovina, J.J., Howell, E., Kobayashi, D.R., and Seki, M.P.
(2001) The transition zone chlorophyll front, a dynamic
global feature defining migration and forage habitat for
marine resources. Progress in Oceanography, 49,
469–83.
Pomilla, C. and Rosenbaum, H.C. (2005) Against the current: an inter-oceanic whale migration event. Biology
Letters, 1, 476–79.
Portugal, S.J., Green, J.A., and Butler, P.J. (2007) Annual
changes in body mass and resting metabolism in captive barnacle geese (Branta leucopsis): the importance of
wing moult. Journal of Experimental Biology, 210,
1391–97.
Post, D.M., Taylor, J.P., Kitchell, J.F., Olson, M.H., Schindler,
D.E., and. Herwig, B.R. (1998) The role of migratory
waterfowl as nutrient vectors in a managed wetland.
Conservation Biology, 12, 910–20.
Potts, W.K. (1984) The chorus-line hypothesis of manoeuvre coordination in avian flocks. Nature, 309, 344–45.
Prins, H.H.T. (1989) East African grazing lands: overgrazed or stably degraded. In W.D. Verwey (ed.) Nature
Management and Sustainable Development, IOS:
Amsterdam/Tokyo, pp. 281–306.
Prins, H.H.T. (1992) The pastoral road to extinction—competition between wildlife and traditional pastoralism in
East Africa. Environmental Conservation, 19, 117–23.
Pulido, F. (2007) The genetics and evolution of avian
migration. Bioscience, 57, 165–74.
Pulido, F. and Berthold, P. (2003) The quantitative genetic
analyses of migratory behavior. In P. Berthold and
E. Gwinner (eds) Avian migration, pp. 53–77. Springer:
Heidelberg.
Pulido, F., Berthold, P., and van Noordwijk, A.J. (1996)
Frequency of migrants and migratory activity are genetically correlated in a bird population: evolutionary
implications. Proceedings of the National Academy of
Sciences, 93, 14642–47.
Quinn, T.P. (2005) The Behavior and Ecology of Pacific
Salmon and Trout, American Fisheries Society, Bethesda
in association with University of Washington Press:
Seattle and London.
Quinn, T.P., Hodgson, S., and Peven, C. (2007) Temperature,
flow, and the migration of adult sockeye salmon
(Oncorhynchus nerka) in the Columbia River. Can. Journal
of Fisheries and Aquatic Sciences, 54, 1349–60.
Quinn, T.P., Unwin, M.J., and Kinnison, M.T. (2000)
Evolution of temporal isolation in the wild: genetic
divergence in timing of migration and breeding by
introduced Chinook salmon populations. Evolution, 54,
4, 1372–85.
Racey, G., Harris, A., Gerrish, L., Armstrong, E., McNicol,
J., and Baker, J. (1999) Forest Management Guidelines
for the Conservation of Woodland Caribou: a Landscape
Approach. MS draft. Ontario Ministry of Natural
Resources: Thunder Bay, Ontario.
Radakov, D.V. (1973), Schooling in the Ecology of Fish,
J. Wiley & Sons: New York.
Raffalovich, L.E., Deane, G.D., Armstrong, D., and Tsao,
H.S. (2008) Model selection procedures in social research:
Monte-Carlo simulation results. Journal of Applied
Statistics, 35, 1093–114.
Raine, A.F. (2007) The International Impact of Hunting
and Trapping in the Maltese Islands, BirdLife Malta
(internal report).
Ramenofsky, M. and Wingfield, J.C. (2007) Regulation of
migration. Bioscience, 57, 135–44.
Ramenofsky, M., Savard, R., and Greenwood, M.R.C.
(1999) Seasonal and diet transitions in physiology and
behavior in the migratory dark-eyed junco. Comparative
Biochemistry and Physiology A, 122, 385–97.
Rankin, MA. and Burchsted, J.C.A. (1992) The cost of
migration in insects. Annual Review of Entomology, 37,
533–59.
Rannestad, O.T., Danielsen, T., Moe, S.R., and Stokke, S.
(2006) Adjacent pastoral areas support higher densities
of wild ungulates during the wet season than the Lake
Mburo National Park in Uganda. Journal of Tropical
Ecology, 22, 675–83.
Rappole, J.H. (2003) An integrative framework for understanding the origin and evolution of avian migration.
Journal of Avian Biology, 34, 124–28.
Rappole, J.H. and Warner, D.W. (1976) Relationships
between behavior, physiology and weather in avian
transients at a migration stopover site. Oecologia, 26,
193–212.
Rappole, J.H. and Jones, P. (2002) Evolution of Old and
New World migration systems. Ardea, 90, 525–37.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Ratikainen, I.L., Gill, J.A., Gunnarsson, T.G., Sutherland,
W.J., and Kokko, H. (2007) When density-dependence is
not instantaneous: theoretical developments and management implications. Ecology Letters, 10, 1–15.
Rautenstrauch, K.R. and Krausman, P.R. (1989) Influence
of water availability and rainfall on movements of
desert mule deer. Journal of Mammalogy, 70, 197–201.
Rayfield, B., James, P., Fall, A., and Fortin, M.-J. (2008)
Comparing static versus dynamic protected areas in
dynamic boreal ecosystems. Biological Conservation,
141, 438–49.
Reed, J. M. (2004) Recognition behavior based problems in
species conservation. Annales Zoologici Fennici, 41,
859–877.
Rees, W.G., Stammler, F.M., Danks, F.S., and Vitebksy, P.
(2008) Vulnerability of European reindeer husbandry to
global change. Climatic Change, 87, 199–217.
Reynolds, D.R. and Riley, J.R. (1997) Flight Behaviour and
Migration of Insect Pests: Radar Studies in Developing
Countries, NRI Bulletin 71. Natural Resources Institute:
Chatham, UK.
Reynolds, D.R. and Riley, J.R. (2002) Remote-sensing, telemetric and computer-based technologies for investigating insect movement: a survey of existing and
potential techniques. Computers and Electronics in
Agriculture, 35, 271–307.
Reynolds, D.R., Smith, A.D., and Chapman, J.W. (2008) A
radar study of emigratory flight and layer formation by
insects at dawn over southern Britain. Bulletin of
Entomological Research, 98, 35–52.
Richardson, P.B., Broderick, A.C., Campbell, L.M., Godley,
B.J., and Ranger, S. (2006) Marine turtle fisheries in the
UK Overseas Territories of the Caribbean: domestic legislation and the requirements of multilateral agreements. Journal of International Wildlife Law and Policy,
9, 223–46.
Richardson, W.J. (1978) Timing and amount of bird
migration in relation to weather: A review. Oikos, 30,
224–72.
Richardson, W.J. (1979) Southeastward shorebird migration over Nova Scotia and New Brunswick in autumn—
Radar study. Canadian Journal of Zoology/Revue
canadienne de zoologie, 57, 107–24.
Richardson, W.J. (1990) Timing and amount of bird migration in relation to weather: updated review. In E Gwinner
(ed.) Bird Migration: Physiology and Ecophysiology,
pp. 78–101, Springer: Berlin, Heidelberg.
Ricker, W. (1954) Stock and recruitment. Journal of the
Fisheries Research Board of Canada, 11, 559–623.
Ricklefs, R.E. (2002) Splendid isolation: historical ecology
of the South American passerine fauna. Journal of Avian
Biology, 33, 207–11.
245
Roberge, J.M. and Angelstam, P. (2004) Usefulness of the
umbrella species concept as a conservation tool.
Conservation Biology, 18, 1, 76–85.
Robinson, D.W., Bowlin, M.S., Bisson, I., Shamoun-Baranes,
J., Thorup, K., Diehl, R., Kunz, T., Mabey, S., and Winkler,
D.W. (2010) Integrating concepts and technologies to
advance the study of bird migration. Frontiers in Ecology
and the Environment, doi:10.1890/080179.
Robinson, R.A., Learmouth, J.A., Hutson, A.M., MacLeod,
C.D., Sparks, T.H., Leech, D.I., Pierce, G.J., Rehfische,
M.M., and Crick, H.Q.P. (2005) Climate Change and
Migratory Species, British Trust for Ornithology:
Thetford, UK.
Robinson R.A., Crick H.Q.P., Learmonth J.A., Maclean
I.M.D., Thomas C.D., Bairlein F., Forchhammer M.C.,
Francis C.M., Gill J.A., Godley B.J., Harwood J., Hays
GC., Huntley B., Hutson A.M., Pierce G.J., Rehfisch M.M.,
Sims D.W., Santos B.M., Sparks T.H., Stroud D.A., and
Visser M.E. (2008) Travelling through a warming world:
climate change and migratory species, Endangered
Species Research, 7, 87–99.
Robinson, S. and Milner-Gulland, E.J. (2003) Contraction
of livestock mobility resulting from state far reorganisation. In C.K. Kerven, (ed.) Prospects for Pastoralism in
Kazakhstan and Turkmenistan: From State Farms to
Private Flocks, RoutledgeCurzon: London.
Robinson, S. and Milner-Gulland, E.J. (2003) Political
change and factors limiting numbers of wild and
domestic ungulates in Kazakhstan. Human Ecology,
31, 87–110.
Robinson, S., Milner-Gulland, E.J., and Alimaev, I. (2003)
Rangeland degradation in Kazakhstan during the Soviet
era: re-examining the evidence. Journal of Arid
Environments, 53, 419–39.
Robinson, T. and Minton, C. (1989) The enigmatic banded
stilt. Birds International, 1, 72–85.
Roed, K., Flagstad, Ø., Nieminen, M., Holand, Ø., Dwyer,
M., Røv, N., and Vilà, C. (2008) Genetic analyses reveal
independent domestication origins of Eurasian Reindeer.
Proceedings of the Royal Society, 275, 1849–55.
Roff, D.A. (1990) The evolution of flightlessness in insects.
Ecological Monographs, 60, 389–421.
Roff, D.A. and Fairbairn, D.J. (2007) The evolution and
genetics of migration in insects. Bioscience, 57, 155–64.
Romanczuk, P., Couzin, I.D., and Schimansky-Geier, L.
(2009) Collective motion due to individual escape and
pursuit response. Physical Review Letters, 102, 4.
Ropert-Coudert, Y. and Wilson, R.P. (2005) Trends and perspectives in animal-attached remote sensing. Frontiers
in Ecology and the Environment, 3, 437–44.
Rosenzweig, C., Iglesias, A., Yang, X.B., Epstein, P.R., and
Chivian, E. (2001) Climate change and extreme weather
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
246
REFERENCES
events: implications for food production, plant diseases,
and pests. Global Change and Human Health, 2, 2, 90–104.
Roshier, D., Asmus, M., and Klaassen, M. (2008) What
drives long-distance movements in the nomadic grey
teal Anas gracilis in Australia? Ibis, 150, 474–84.
Royle, J.A. and R.M. Dorazio. (2008) Hierarchical Modeling
and Inference in Ecology: The Analysis of Data from
Populations, Metapopulations and Communities,
Academic Press, Elsevier: London.
Rubolini, D., Møller, A.P., Rainio, K. and Lehikoinen, E.
(2007) Intraspecific consistency and geographic variability in temporal trends of spring migration phenology
among European bird species. Climate Research, 35,
135–46.
Rudnick, J.A., Katzner, T.E., Bragin, E.A., and deWoody,
A.J. (2008) A non-invasive genetic evaluation of population size, natal philopatry, and roosting behavior of nonbreeding eastern imperial eagles (Aquila heliaca) in
central Asia. Conservation Genetics, 9, 667–76.
Ruess, R.W. and Seagle, S.W. (1994) Landscape patterns in
soil microbial processes in the Serengeti National Park,
Tanzania. Ecology, 75, 892–904.
Rutz, C. and Hays, G.C. (2009) New frontiers in biologging science. Biology Letters, 5, 289–92.
Sabo, J.L. and Power, M.E. (2002) River-watershed
exchange: Effects of riverine subsidies on riparian lizards and their terrestrial prey. Ecology, 83, 1860–69.
Sadovy, Y. and Domeier, M. (2005) Are aggregation-fisheries sustainable? Reef fish fisheries as a case study. Coral
Reefs, 24, 2, 254–62.
Saiga News. (2009) Updates, 8, 2–7.
Saino, N. and Ambrosini, R. (2008) Climatic connectivity
between Africa and Europe may serve as a basis for
phenotypic adjustment of migration schedules of transSaharan migratory birds. Global Change Biology, 14,
250–63.
Sala, E., Ballesteros, E., and Starr, R.M. (2001) Rapid decline
of Nassau grouper spawning aggregations in Belize:
fishery management and conservation needs. Fisheries,
26, 23–30.
Salewski, V. and Bruderer, B. (2007) The evolution of bird
migration—a synthesis. Naturwissenschaften, 94, 268–79.
Salewski, V. and Jones, P. (2006) Palearctic passerines in
Afrotropical environments: a review. Journal of
Ornithology, 147, 192–201.
Sanderson, F.J., Donald, P.F., Pain, D.J., Burfield, I.J., and
van Bommel, F.P.J. (2006) Long-term population declines
in Afro-Palearctic migrant birds. Biological Conservation,
131, 93–105.
Sapir, N. (2009) The effects of weather on Bee-eater (Merops
apiaster) migration, PhD thesis, The Hebrew University
of Jerusalem.
Sapir, N., Abramsky, Z., Shochat, E., and Izhaki, I.
(2004a) Scale-dependent habitat selection in migratory
frugivorous passerines. Naturwissenschaften, 91, 544–47.
Sapir, N., Tsurim, I., Gal, B., and Abramsky, Z. (2004b) The
effect of water availability on fuel deposition of two staging Sylvia warblers. Journal of Avian Biology, 35, 25–32.
Sato, K., Watanuki, Y., Takahashi, A., et al. (2007) Stroke
frequency, but not swimming speed, is related to body
size in free-ranging seabirds, pinnipeds and cetaceans.
Proceedings of the Royal Society B-Biological Sciences,
274, 471–77.
Satterthwaite, W.H., Beakes, M.P., Collins, E., Swank, D.R.,
Merz, J.E., Titus, R.G., Sogard, S.M., and Mangel, M.
(2009) Steelhead life history on California’s central coast:
insights from a state dependent model. Transactions of
the American Fisheries Society, 138, 532–48.
Satterthwaite, W.H., Beakes, M.P., Collins, E., Swank, D.R.,
Merz, J.E., Titus, R.G., Sogard, S.M., and Mangel, M.
(2010) State-dependent life history models in a changing (and regulated) environment: steelhead in the
California Central Valley. Evolutionary Applications,
3, 221– 43.
Sauerbrei, W., Hollander, N., and Buchholz, A. (2008)
Investigation about a screening step in model selection.
Statistics and Computing, 18, 195–208.
Sawyer, H., Kauffman, M.J., Nielson, R.M., and Horne, J.S.
(2009) Identifying and prioritizing ungulate migration
routes for landscape-level conservation. Ecological
Applications, 19, 2016–25.
SCFC (Siberian Crane Flyway Coordination) (Undated
a)Reintroduction: Hang-glider Migration, [Online],
Available at: http://www.sibeflyway.org/ReintroductionHang-glider%20Mig-web.html. [August 2009].
SCFC (Siberian Crane Flyway Coordination) (Undated b)
Reintroduction. [Online], Available at: http://www.
sibeflyway.org/Reintroduction-Russia-web.html#kurn.
[August 2009].
Schaefer, J.A. (2003) Long-term range recession and the
persistence of caribou in the taiga. Conservation Biology,
17, 1435–39.
Schaefer, J.A., Bergman, C.M., and Luttich, S.N. (2000) Site
fidelity of female caribou at multiple spatial scales.
Landscape Ecology, 15, 731–39.
Schaefer, J.A., Veitch, A.M., Harrington, F.H., Brown, W.K.,
Theberge, J.B., and Luttich, S.N. (2001) Fuzzy structure
and spatial dynamics of a declining woodland caribou
population. Oecologia, 126, 507–14.
Schaub, M., Jenni, L., and Bairlein, F. (2008) Fuel stores,
fuel accumulation, and the decision to depart from a
migration stopover site. Behavioral Ecology, 19, 657–66.
Schaub, M., Pradel, R., and Lebreton, J.-D. (2004) Is the
reintroduced white stork (Ciconia ciconia) population
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
in Switzerland self-sustainable? Biological Conservation,
119, 105–14.
Schick, R.S., S.R. Loarie, F. Colchero, B.D. Best, A. Boustany,
D.A. Conde, P.N. Halpin, L.N. Joppa, C.M. McClellan,
and J.S. Clark (2008) Understanding movement data
and movement processes: current and emerging directions. Ecology Letters, 11, 1338–50.
Schlecht, E., P. Hiernaux, and M.D. Turner (2001) Mobilité
régionale du bétail: nécessité et alternatives? In E.
Tielkes, E. Schlecht, and P. Hiernaux (eds) Elevage et
gestion de parcours au Sahel, implications pour le
développement, Verlag Grauer: Beuren-Stuttgart.
Schlichting, C.D. and Pigliucci, M. (1998) Phenotypic evolution: a reaction norm perspective. Sinauer Associates,
Inc., Sunderland.
Schmaljohann, H. and Dierschke, V. (2005) Optimal bird
migration and predation risk: a field experiment with
northern wheatears Oenanthe oenanthe. Journal of
Animal Ecology 74, 131–38.
Schmidt, K.A. (2004) Site fidelity in temporally correlated
environments enhances population persistence. Ecology
Letters, 7, 176–84.
Schmidt-Nielsen, K. (1972) Locomotion: energetic cost of
swimming, flying and running. Science, 177, 222–28.
Schmidt-Nielsen, K. (1975) Animal Physiology. Cambridge
University Press: Cambridge.
Schmidt-Wellenburg, C.A., Engel, S., and Visser, G.H.
(2008) Energy expenditure during flight in relation to
body mass: effects of natural increases in mass and artificial load in Rose Coloured Starlings. Journal of
Comparative Physiology B-Biochemical Systemic and
Environmental Physiology, 178, 767–77.
Schmitz, J. (1986) L’état géomètre: les leydi des Peul du
Fuuta Tooro (Sénégal) et du Maasina (Mali). Cahiers
d’Etudes Africaines, 26, 3, 349–94.
Schoenecker, K.A., Singer, F.J., Zeigenfuss, L.C., Binkley,
D., and Menezes, R.S.C. (2004) Effects of elk herbivory
on vegetation and nitrogen processes. Journal of Wildlife
Management, 68, 837–49.
Schuman, J.R. (1995) Environmental considerations for
assessing dam removal alternatives for river restoration.
Regulated Rivers: Research and Management, 11, 249–61.
Scoones, I. (1995) Exploiting heterogeneity: habitat use by
cattle in dryland Zimbabwe. Journal of Arid
Environments, 29, 221–37.
Seabloom, E.W., Dobson, A.P., and Stoms, D.M. (2002)
Extinction rates under nonrandom patterns of habitat
loss. Proceedings of the National Academy of Sciences,
99, 17, 11229–34.
Seagle, S.W. (2003) Can ungulates foraging in a multipleuse landscape alter forest nitrogen budgets? Oikos 103,
230–34.
247
Seagle, S.W., S.J. McNaughton, and R.W. Ruess (1992)
Simulated effects of grazing on soil nitrogen and mineralization in contrasting Serengeti grasslands. Ecology,
73, 1105–23.
Secundus, C.P. (77) Historia Naturalis.
Seeley, T.D. (1985), Honeybee ecology, Princeton University
Press: Princeton.
Seiler, A., Cedarlund, G, Jernelid, H, Grängstedt P., and
Ringaby, E. (2003) The barrier effect of highway E4 on
migratory moose (Alces alces) in the High Coast area,
Sweden. Proceedings of the IENE conference on Habitat
fragmentation due to transport infrastructure, Brussels,
13–14 November 2003, [Online], Available at: http://
w w w. g r i m s o . s l u . s e / re s e a rc h / i n f r a s t r u c t u re /
Documents/HighCoast_IENE2003.pdf. [March 2009]
Senft, R.L., Coughenour, M.B., Bailey, D.W., Rittenhouse,
L.R., Sala, O.E., and Swift, D.M. (1987) Large herbivore
foraging and ecological hierarchies: landscape ecology
can enhance traditional foraging theory. BioScience, 37,
11, 789–99.
Serneels, S. and Lambin, E. (2001) Impact of land-use
change on the wildebeest migration in the northern part
of the Serengeti-Mara ecosystem. Journal of Biogeography, 28, 391–407.
Shahgedanova, M. (ed.) (2003) The Physical Geography of
Northern Eurasia, Oxford University Press: Oxford.
Shamoun-Baranes, J., Liechti, O., Yom-Tov, Y., and Leshem,
Y. (2003a) Using a convection model to predict altitudes
of white stork migration over central Israel. BoundaryLayer Meteorology, 107, 673–81.
Shamoun-Baranes, J., Leshem, Y., Yom-Tov, Y., and
Liechti, O. (2003b) Differential use of thermal convection by soaring birds over central Israel. Condor, 105,
208–18.
Shannon, H., Young, G., Yates, M., Fuller, M., and Seegar,
W. (2002) American white pelican soaring flight times
and altitudes relative to changes in thermal depth and
intensity. Condor, 104, 679–83.
Shea, R.E., Nelson, H.K., Gillette, L.N., King, J.G., and Weaver,
D.K. (2002) Restoration of Trumpeter Swans in North
America: A Century of Progress and Challenges.
Waterbirds: The International Journal of Waterbird Biology,
25, Special Publication 1, Proceedings of the Fourth
International Swan Symposium, 2001, pp. 296–300.
Shepherd, B. and Whittington, J. (2006) Response of
wolves to corridor restoration and human use management. Ecology and Society, 11, 2, 1. [Online], Available:
http://www.ecologyandsociety.org/vol11/iss2/
art1/[November 2009].
Sherrill-Mix, S.A., James, M.C., and Myers, R.A. (2008)
Migration cues and timing in leatherback sea turtles.
Behavioral Ecology, 19, 2, 231–236.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
248
REFERENCES
Shields, E. J. and Testa, A. M. (1999) Fall migratory flight
initiation of the potato leafhopper, Empoasca fabae
(Homoptera: Cicadellidae): observations in the lower
atmosphere using remote piloted vehicles. Agricultural
and Forest Meteorology, 97, 317–30.
Shillinger, G.L., Palacios, D.M., Bailey, H., Bograd, S.J.,
Swithenbank, A.M., Gaspar, P., Wallace, B.P., Spotila,
J.R., Paladino, F.V., Piedra, R., Eckert, S.A and, Block,
B.A. (2008) Persistent leatherback turtle migrations
present opportunities for conservation. Public Library
of Science, Biology, 6, 7, 1408–16.
Shine, R. and Brown, G.P. (2008) Adapting to the unpredictable: reproductive biology of vertebrates in the
Australian wet-dry tropics. Philosophical Transactions
of the Royal Society of London, Series B, 363, 363–73.
Sibert, J.R. and Fournier, D.A. (2001) Possible Models for
combining tracking data with conventional tagging
data. Symposium on Tagging and Tracking Marine Fish
with Electronic Devices, February 7–11 2000, Honolulu.
In J. R. Sibert and J. L. Nielsen (eds) Electronic Tagging
and Tracking in Marine Fisheries, Kluwer Academic
Press.
Sih, A., Bell, A.M., Johnson, J.C., and Ziemba, R.E. (2004)
Behavioral syndromes: an integrative overview. The
Quarterly Review of Biology, 79, 241–77.
Sillett, T.S. and Holmes, R.T. (2002) Variation in survivorship of a migratory songbird throughout its annual
cycle. Journal of Animal Ecology, 71, 296–308.
Simons, A.M. (2004) Many wrongs: the advantage of
group navigation. Trends in Ecology and Evolution 19,
453–55.
Simpson, S.J. and Miller, G.A. (2007) Maternal effects on
phase characteristics in the desert locust, Schistocerca
gregaria: A review of current understanding. Journal of
Insect Physiology, 53, 869–76.
Simpson, S.J. and Sword, G.A. (2008) Locusts. Current
Biology, 18, 364–66.
Simpson, S.J. and Sword, G.A. (2009) Phase polyphenism
in locusts: Mechanisms, population consequences,
adaptive significance and evolution. In D. Whitman and
T. Ananthakrishnan (eds) Phenotypic Plasticity of
Insects: Mechanisms and Consequences, pp 147–90,
Science Publishers Inc.: Plymouth.
Simpson, S.J., Despland, E., Hagele, B.F., and Dodgson, T.
(2001) Gregarious behavior in desert locusts is evoked
by touching their back legs. Proceedings of the National
Academy of Sciences of the United States of America,
98, 3895–97.
Simpson, S.J., Sword, G.A., Lorch, P.D., and Couzin, I.
D. (2006) Cannibal crickets on a forced march for protein and salt. Proceedings of the National Academy of
Sciences of the United States of America, 103,
4152–56.
Sims, D.W., Southall, E.J., Humphries, N.E., et al. (2008)
Scaling laws of marine predator search behaviour.
Nature, 451, 1098–1102.
Sinclair, A.R.E. (1977) The African Buffalo: a Study of
Resource Limitation of Populations, University of
Chicago Press, Chicago.
Sinclair, A.R.E. (1978) Factors affecting food supply and
breeding season of resident birds and movements of
palaearctic migrants in a tropical African savanna. Ibis,
120, 480–97.
Sinclair, A.R.E. (1979) The Serengeti environment, pp.
31–45 in A.R.E. Sinclair (ed.) Serengeti—Dynamics of an
Ecosystem. University of Chicago Press, Chicago.
Sinclair, A.R.E. (1983) The function of distance movement
in vertebrates. In I.R. Swingland and P.J. Greenwood
(eds) The Ecology of Animal Movement, pp. 248–58,
Clarendon Press: Oxford.
Sinclair, A.R.E. (1985) Does interspecific competition or
predation shape the African ungulate community.
Journal of Animal Ecology 54, 899–918.
Sinclair, A.R.E., Dublin, H., and Borner, M. (1985)
Population regulation of Serengeti wildebeest—a test of
the food hypothesis. Oecologia, 65, 266–68.
Sinclair, A.R.E., Mduma, S., and Brashares, J.S. (2003)
Patterns of predation in a diverse predator-prey system.
Nature, 425, 288–90.
Sinclair, A.R.E., Mduma, S.A.R., Hopcraft, J.G.C., Fryxell,
J.M., Hilborn, R., and Thirgood, S. (2007) Long-term
ecosystem dynamics in the Serengeti: Lessons for conservation. Conservation Biology, 21, 580–90.
Singh, N., Milner-Gulland, E.J. (in press) Conserving a
moving target: Planning protection for a migratory species as its distribution changes. Journal of Applied Ecology
Singh, N.J., Grachev, Iu.A., Bekenov, A.B., and MilnerGulland, E.J. (2010) Tracking greenery in Central Asia:
The migration of the saiga antelope. Diversity and
Distributions 16, 663–75.
Skelly, D.K. and Werner, E.E. (1990) Behavioral and lifehistorical responses of larval American toads to an odonate predator. Ecology, 71, 2313–22.
Sladen, W.J.L. and Olsen, G.H. (2005) Teaching geese,
swans and cranes pre-selected migration routes using
ultralight aircraft, 1990–2004—looking into the future.
In M.H. Linck and R.E. Shea (eds) Selected papers of the
twentieth trumpeter swan society conference—Trumpeter Swan Restoration: Exploration and Challenges,
20–22 October 2005, Council Bluffs, Iowa, pp. 53–54.
Sladen, W.J.L., Lishman, W.A., Ellis, D.H., Shire, G.G., and
Rininger, D.L. (2002) Teaching migration routes to canada
geese and trumpeter swans using ultralight aircraft, 1990–
2001. Waterbirds, The International Journal of Waterbird
Biology, 25, Special Publication 1: Proceedings of the
Fourth International Swan Symposium, 2001, pp. 132–37.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Slotte, A. and Fiksen, Ø. (2000) State-dependent spawning
migration in Norwegian spring-spawning herring.
Journal of Fish Biology, 56, 138–62.
Small, C.J. (2005) Regional Fisheries Management Organisations: Their Duties and Performance in Reducing
Bycatch of Albatrosses and Other Species, BirdLife
International: Cambridge, UK.
Smith, R.J. and Moore, F.R. (2003) Arrival fat and reproductive performance in a long-distance passerine
migrant. Oecologia, 134, 325–31.
Smith, R.J. and Moore, F.R. (2005) Arrival timing and seasonal reproductive performance in a long-distance
migratory landbird. Behavioral Ecology and
Sociobiology, 57, 231–39.
Smith, S.B. and McWilliams, S.R. (2010) Patterns of fuel
use and storage in migrating passerines in relation to
fruit resources at autumn stopover sites. Auk, 127,
108–18.
Smuts, G. L. (1978) Interrelations between Predators, Prey,
and Their Environment. Bioscience, 28, 316–20.
Sniegowski, P.D., Ketterson, E.D., and Nolan, V. (1988)
Can experience alter the avian annual cycle—Results of
migration experiments with Indigo Buntings. Ethology,
79, 333–41.
Snow, D. and Perrins, C. (1998) The Complete Birds of the
Western Palearctic on CD-ROM, Oxford University
Press: Oxford, England.
Sokolov, V.E. and Zhirnov, L.V. (eds) (1998) The Saiga
Antelope; Phylogeny, Systematics, Ecology, Conservation and Use, Russian Academy of Sciences:
Moscow.
Solbreck, C. (1978) Migration, diapause, and direct development as alternative life histories in a seed bug,
Neacoryphus bicrucis. In H. Dingle (ed.) Evolution of
Insect Migration and Diapause, pp.195–217, Springer:
New York.
Sorokin, A., Shilina, A., Ermakov, A., and Markin, Y. (2002)
Hang-glider Migration: Flight of Hope, [Online],
Available at: http://www.sibeflyway.org/Reintroduction-Flight-of-Hope-Project-web.html.
[August
2009].
Soulé, M.E. (1985) What is conservation biology?
BioScience, 35, 727–34.
Soulé, M.E. and Kohm, K. (1989) Research Priorities in
Conservation Biology, Island Press, Washington, DC.
Southwood, A. and Avens, L. (2010) Physiological, behavioural, and ecological aspects of migration in reptiles.
Journal of Comparative Physiology Part B, 180, 1–23.
Southwood, A, Fritsches, K, Brill, R, and Swimmer, Y.
(2008) Sound, chemical, and light detection in sea turtles
and pelagic fishes: sensory-based approaches to bycatch
reduction in longline fisheries. Endangered Species
Research, 5, 225–38.
249
Southwood, T.R.E. (1962) Migration of terrestrial arthropods in relation to habitat. Biological Reviews, 37,
171–214.
Southwood, T.R.E. (1977) Habitat, the templet for ecological strategies? Journal of Animal Ecology, 46, 337–65.
Southwood, T.R.E. (1981) Ecological aspects of insect
migration. In D.J. Aidley (ed.) Animal Migration,
pp. 197–208, Cambridge University Press: Cambridge.
Sparks, T.H., Dennis, R.L.H., Croxton, P.J. and Cade, M.,
(2007) Increased migration of Lepidoptera linked to
climate change. European Journal of Entomology, 104,
139–43.
Spiegelhalter, D. and Rice, K. (2009) Bayesian statistics.
Scholarpedia, 4, 8, 5230.
Spotila, J., Reina, R.D., Steyermark, A.C., Plotkin, P.T., and
Spotila, F.V. (2000) Pacific leatherback turtles face extinction. Nature, 405, 529–31.
Srygley, R.B. and Dudley, R. (2008) Optimal strategies for
insects migrating in the flight boundary layer: mechanisms and consequences. Integrative and Comparative
Biology, 48, 119–33.
Stammler, F. (2005) Reindeer Nomads Meet the Market:
Culture, Property and Globalisation at the End of the
Land. Vol. 6. Halle Studies in the Anthropology of
Eurasia. Münster: Lit publishers.
Stammler, F.(2009) Mobile phone revolution in the tundra? Technological change among Russian reindeer
nomads. In A. Ventsel (ed.) Generation P in the Tundra,
Folklore, 41, 47–78, Estonian Literary Museum:
Talinn.
Stammler, F. and Beach, H. (eds) (2006) People and
Reindeer on the Move. Special Issue of the journal
Nomadic Peoples, 10, 2, Berghahn Publishers: Oxford.
Stammler, F. and Peskov, V. (2008) Building a ‘Culture of
dialogue’ among stakeholders in North-West Russian
oil extraction. Europe-Asia Studies, 60, 5, 831–849.
Standen, E.M., Hinch, S.G., and Rand, P.S. (2004)
Influence of river speed on path selection by migrating adult sockeye salmon (Oncorhynchus nerka)
Canadian Journal of Fisheries and Aquatic Sciences,
61, 905–12.
Stanley, E.H. and Doyle, M.W. (2003) Trading off: the ecological effects of dam removal. Frontiers in Ecology and
the Environment, 1, 15–22.
Stanley, E.H., Catalano, M.J., Mercado-Silva, N., and Orr,
C.H. (2007) Short communication: effects of dam
removal on brook trout in a Wisconsin stream. River
Research and Applications, 23, 792–98.
Stapp, P. and Polis, G.A. (2003) Marine resources subsidize
insular rodent populations in the Gulf of California,
Mexico. Oecologia, 134, 496–504.
Stefansson, S.O., Björnsson, B.Th., and McCormick, S.D.
(2008) Smoltification. In R. N. Finn and B. K. Kapo (eds)
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
250
REFERENCES
Fish Larval Physiology, pp. 639–81, Science Publishers:
Enfield NH, USA.
Stefansson, S.O., Nilsen, T.O., Ebbesson, L.O.E., Wargelius,
A., Madsen, S.S., BjÖrnsson, B.T., and McCormick, S.D.
(2007) Molecular mechanisms of continuous light inhibition of Atlantic salmon parr-smolt transformation.
Aquaculture, 273, 235–45.
Stenning, D.J. (1957) Transhumance, migratory drift,
migration: patterns of pastoral Fulani nomadism. Journal
of the Royal Anthropological Institute, 87, 57–73.
Stephens, D.M. and Krebs, J.R. (1986) Foraging Theory,
Princeton University Press, Princeton, New Jersey.
Stephens, D.W., Brown, J.S., and Ydenberg, R.C. (2007)
Foraging Behavior and Ecology. University of Chicago
Press: Chicago.
Stige, L.C., Chan, K.-S., Zhang, Z., Frank, D., and Stenseth,
N.C. (2007) Thousand-year-long Chinese time series
reveals climatic forcing of decadal locust dynamics.
Proceedings of the National Academy of Sciences USA,
104, 16188–93.
Stoddard, P.K., Marsden, J.E., and Williams, T.C. (1983)
Computer simulation of autumnal bird migration over the
western North Atlantic. Animal Behaviour, 31, 173–80.
Strandberg, R. (2008) Migration strategies of raptors:
Spatio-temporal adaptations and constraints in travelling and foraging. Ph.D. thesis, Lund University: Lund.
Strandberg, R. and Alerstam, T. (2007) The strategy of flyand-forage migration, illustrated for the osprey (Pandion
haliaetus). Behavioral Ecology and Sociobiology, 61,
1865–75.
Strandberg, R., Klaassen, R.H.G., Hake, M., and
Alerstam, T. (2009) How hazardous is the Sahara
Desert crossing for migratory birds? Indications
from satellite tracking of raptors. Biology Letters,
doi:10.1098/rsbl.2009.0785.
Strandberg, R., Klaassen, R.H.G., Hake, M., Olofsson, P.,
and Alerstam, T. (2009) Converging migration routes of
Eurasian hobbies Falco subbuteo crossing the African
equatorial rain forest. Proceedings of the Royal Society
B-Biological Sciences, 276, 727–33.
Strindberg, S. and Buckland, S.T. (2004) Zigzag Survey
Designs in Line Transect Sampling. Journal of Agricultural,
Biological, and Environmental Statistics, 9, 443–61.
Studds, C.E. and Marra, P.P. (2005) Nonbreeding habitat
occupancy and population processes: an upgrade experiment with a migratory bird. Ecology, 86, 2380–85.
Studds, C.E. and Marra, P.P. (2007) Linking fluctuations in
rainfall to nonbreeding season performance in a longdistance migratory bird, Septophaga ruticilla. Climate
Research, 35, 115–22.
Stull, R.B. (1988) An Introduction to Boundary Layer
Meteorology, Kluwer Academic Publishers: Boston.
Stutchbury, B.J.M., Tarof, S.A., Done, T., Gow, E., Kramer,
P.M., Tautin, J., Fox, J.W., and Afanasuev, V. (2009)
Tracking long-distance songbird migration by using
geolocators. Science, 323, 896.
Sunnucks, P. (2000) Efficient genetic markers for population biology. Trends in Ecology and Evolution, 15,
199–203.
Sutherland, W.J. (1983) Aggregation and the ‘Ideal Free’
distribution. Journal of Animal Ecology, 52, 821–28
Sutherland, W.J. (1996) From Individual Behaviour to
Population Ecology, Oxford University Press: Oxford.
Sutherland, W.J. (1998) Evidence for flexibility and constraint in migration systems. Journal of Avian Biology,
29, 441–46.
Sutherland, W.J., Clout, M., Cote, I.M., Daszak, P.,
Depledge, M.H., Fellman, L., Fleishman, E., Garthwaite,
R., Gibbons, D.W., De Lurio, J., Impey, A.J., Lickorish, F.,
Lindenmayer, D., Madgwick, J., Margerison, C.,
Maynard, T., Peck, L.S., Pretty, J., Prior, S., Redford,
K.H., Scharlemann, J.P.W., Spalding, M., and Watkinson,
A.R. (2010) A horizon scan of global conservation issues
for 2010. Trends in Ecology and Evolution, 25, 1–7.
Svedang, H. and Wickstrom, H. (1997) Low fat contents in
female silver eels: indications of insufficient energetic
stores for migration and gonadal development. Journal
of Fish Biology, 50, 475–86.
Switzer, P.V. (1997) Past reproductive success affects future
habitat selection. Behavioral Ecology and Sociobiology,
40, 307–12.
Sword, G.A. (2003) To be or not to be a locust? A comparative analysis of behavioral phase change in nymphs of
Schistocerca americana and S. gregaria. Journal of Insect
Physiology, 47, 709–17.
Sword, G.A., Lorch, P.D., and Gwynne, D.T. (2005) Migratory bands give crickets protection. Nature, 433, 703.
Takakura, H. (2002) An institutionalized human-animal
relationship and the aftermath: the reproductive process of horse-bands and husbandry in Northern Yakutia,
Siberia. Human Ecology, 30, 1–19.
Takakura, H. (2004) Gathering and Releasing Animals:
Reindeer herd control activities of the indigenous peoples of the Verkhoyansky Region, Siberia. Bulletin of the
National Museum of Ethnology, 29, 1, 43–70.
Takimoto, G., Iwata, T., and Murakami, M. (2002) Seasonal
subsidy stabilizes food web dynamics: Balance in a heterogeneous landscape. Ecological Research, 17, 433–39.
Talbot, L.M. and Talbot M.H., (1963) The wildebeest in
western Masailand, East Africa. Wildlife Monographs,
12, 1–88.
Taylor, C.M. and Norris, D.R. (2007) Predicting conditions
for migration: effects of density dependence and habitat
quality. Biology Letters, 3, 280–83.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Taylor, L. R. (1974) Insect migration, flight periodicity and
boundary-layer. Journal of Animal Ecology, 43, 225ff.
Taylor, L.R. (1986) Synoptic ecology, migration of the second kind, and the Rothamssted Insect Survey.
Presidential address. Journal of Animal Ecology, 55,
1–38.
Teo, S.L.H., Boustany, A., Dewar, H., et al. (2007) Annual
migrations, diving behavior, and thermal biology of
Atlantic bluefin tuna, Thunnus thynnus, on their Gulf of
Mexico breeding grounds. Marine Biology, 151, 1–18.
Terborgh, J. and van Schaik, C. (2002) Why the world
needs parks. In J. Terborgh, C. van Schaik, L. Davenport,
and M. Rao (eds) Making Parks Work: Strategies for
Preserving Tropical Nature, pp. 3–14, Island Press:
Washington, DC.
Tevis, L., Jr. and Newell, I.M. (1962) Studies on the biology
and seasonal cycle of the Giant Red Velvet Mite,
Dinothrombium pandorae (Acari, Trombidiidae). Ecology,
43, 497–505.
Thirgood, S., Mosser, A., Tham, S., Hopcraft, G.,
Mwangomo, E., Mlengeya, T., Kilewo, M., Fryxell, J.,
Sinclair, A.R.E., and Borner, M. (2004) Can parks protect
migratory ungulates? The case of the Serengeti wildebeest. Animal Conservation, 7, 113–20.
Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M.,
Beaumont, L.J., Collingham, Y.C., Erasmus, B. F. N.,
Ferreira de Siqueira, M., Grainger, A., Hannah, L.,
Hughes, L., Huntley, B., van Jaarsveld, A.S., Midgley,
G.F., Miles, L., Ortega-Huerta, M.A., Peterson, A. T.,
Phillips, O.L., and Williams, S.E. (2004) Extinction risk
from climate change. Nature, 427, 8, 145–48.
Thomson, A.L. (1926) Problems of Bird-Migration,
Witherby: London.
Thorpe, J. E. (1977) Bimodal distribution of length of
juvenile Atlantic salmon (Salmo salar L) under artificial rearing conditions. Journal of Fish Biology, 11,
175–84.
Thorpe, J.E. (1988) Salmon migration. Scientific Progress,
72, 345–70.
Thorup, K., Alerstam, T., Hake, M., and Kjellen, N. (2003)
Bird orientation: compensation for wind drift in migrating raptors is age dependent. Proceedings of the Royal
Society of London Series B-Biological Sciences, 270,
S8–S11.
Thorup, K., Bisson, I. A., Bowlin, M. S., Holland, R. A.,
Wingfield, J. C., Ramenofsky, M., and Wikelski, M. (2007)
Evidence for a navigational map stretching across the
continental US in a migratory songbird. Proceedings of
the National Academy of Sciences USA, 104, 18115–19.
Thrower, F.P., Hard, J.J., and Joyce, J.E. (2004) Genetic
architecture of growth and early life-history transitions
in anadromous and derived freshwater populations of
251
steelhead. Journal of Fish Biology, 65 (Supplement A),
286–310.
Thrush, S.F., Pridmore, R.D., Hewitt J.E., and Cummings,
V.J. (1994) The Importance of Predators on a Sandflat—
Interplay between Seasonal-Changes in Prey Densities
and Predator Effects, Marine Ecology-Progress Series,
107, pp. 211–22.
Tilman, D. (1994) Competition and Biodiversity in Spatially
Structured Habitats. Ecology, 75, 2–16.
Torney, C.N.Z., Neufeld, Z., and Couzin, I.D. (2009)
Context-dependent interaction leads to emergent search
behavior in social aggregates. Proceedings of the
National Academy of Sciences USA, 106, 22055–60.
Torre-Bueno, J.R. (1976) Temperature regulation and heat
dissipation during flight in birds. Journal of Experimental
Biology, 65, 471–82.
Trakimas, G. and Sidaravičius, J. (2008) Road mortality
threatens small northern populations of the European
pond turtle, Emys orbicularis. Acta Herpetologica, 3, 2,
161–66.
Treherne, J.E. and Foster, W.A. (1981) Group transmission
of predator avoidance-behaviour in a marine insect—
the Trafalgar effect. Animal Behaviour, 29, 911–17.
Tremblay, Y., Robinson, P.W., and Costa, D.P. (2009) A parsimonious approach to modeling animal movement
data. PLoS ONE, 4, e4711.
Tremblay, Y., S.A. Shaffer, S.L. Fowler, C.E. Kuhn, B.I.
McDonald, M.J. Weise, C.-A. Bost, H. Weimerskirch, D.
E. Crocker, M.E. Goebel, and D.P. Costa. (2006)
Interpolation of animal tracking data in a fluid environment. Journal of Experimental Biology, 209, 128–40.
Tsurim, I., Sapir, N., Belmaker, J., Shanni, I., Izhaki, I.,
Wojciechowski, M.S., Karasov, W.H., and Pinshow, B.
(2008) Drinking water boosts food intake rate, body
mass increase and fat accumulation in migratory blackcaps (Sylvia atricapilla). Oecologia, 156, 21–30.
Tucker, G. and Goriup, P. (2005) Assessment of the merits
of an instrument under the Convention on Migratory
Species covering migratory raptors in the African–
Eurasian region: status report, DEFRA: Bristol, UK.
Tuisku, T. (2002), Reindeer herding. In Ildikó Lehtinen
(ed.) Siberia: Life on the Taiga and Tundra, pp. 100–107,
National Board of Antiquities: Helsinki.
Turchin, P. (1998) Quantitative Analysis of Movement,
Sinauer Associates, Inc. Publishers: Sunderland, Massachusetts.
Turner, M. D. (1999a) The role of social networks, indefinite boundaries and political bargaining in maintaining the ecological and economic resilience of the
transhumance systems of Sudano-Sahelian West
Africa. In M. Niamir-Fuller (ed.) Managing Mobility
in African Rangelands: the Legitimization of Trans-
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
252
REFERENCES
humance, pp. 97–123, Intermediate Technology Publications: London.
Turner, M.D. (1999b) No space for participation: Pastoralist
narratives and the etiology of park-herder conflict in
southwestern
Niger.
Land
Degradation
and
Development, 10, 4, 343–61.
Turner, M.D. (1999c) Spatial and temporal scaling of grazing impact on the species composition and productivity
of Sahelian annual grasslands. Journal of Arid
Environments, 41, 3, 277–97.
Turner, M.D. (2009) Capital on the move: The changing
relation between livestock and labor in Mali, West
Africa. Geoforum 40, 746–55.
Turner, M.D. and Hiernaux, P. (2008) Changing access to
labor, pastures, and knowledge: The extensification of
grazing management in Sudano-Sahelian West Africa.
Human Ecology, 26, 1, 59–80.
Turner, M. D., Hiernaux, P., and Schlecht, E. (2005) The
distribution of grazing pressure in relation to vegetation
resources in semi-arid West Africa: the role of herding.
Ecosystems, 8, 1432–9840.
USGS Alaska science center bar-tailed godwit updates
website (http://alaska.usgs.gov/science/biology/shorebirds/barg_updates.html), accessed 1 February 2009.
Valenza, J. (1981) Surveillance continue des pâturages
naturels sahéliens. Revue de l’élevage et médecine
vétérinaire des pays tropicaux, 34, 1, 83–100.
van Asch, M. and Visser, M.E. (2007) Phenology of forest caterpillars and their host trees: The importance of synchrony. Annual Review of Entomology, 52, 37–55.
van Bael, S.A., Philpott, S.M., Greenberg, R., Bichier, P., Barber,
N.A,. Mooney, K.A., and Gruner, D.S. (2008) Birds as predators in tropical agroforestry systems. Ecology, 89, 928–34.
van der Jeugd, H.P., Eichhorn, G., Litvins, K.E., Stahl,
J., Larsson, K., van der Graaf, A.J., and Drent, R.H. (2009)
Keeping up with early springs: rapid range expansion
in an avian herbivore incurs a mismatch between reproductive timing and food supply. Global Change Biology,
15, 1057–71.
van Dyne, G.M., Brockington, N.R., Szocs, Z., Duek, J., and
Ribic, C.A. (1980) Large herbivore subsystem. In
Grasslands, Systems, Analysis and Man. International
Biological Programme 19, pp. 269–537, Cambridge
University Press: Cambridge.
van Gils, J. A., Beekman, J. H., Coehoorn, P., Corporaal,
E., Dekkers, T., Klaassen, M., van Kraaij, R., de Leeuw,
R., and de Vries, P.P. (2008) Longer guts and higher food
quality increase energy intake in migratory swans.
Journal of Animal Ecology, 77, 1234–41.
van Gils, J.A., Piersma, T., Dekinga, A., and Battley, P.F.
(2006) Modelling phenotypic flexibility: an optimality
analysis of gizzard size in red knots Calidris canutus.
Ardea, 94, 409–20.
van Ginneken, V.J.T. and van den Thillart, G. (2000)
Physiology—Eel fat stores are enough to reach the
Sargasso. Nature, 403, 156–57.
van Ginneken, V., Antonissen, E., Muller, U.K., Booms,
R., Eding, E., Verreth, J., and van den Thillart, G. (2005)
Eel migration to the Sargasso: remarkably high swimming efficiency and low energy costs. Journal of
Experimental Biology, 208, 1329–35.
van Moorter, B., Visscher, D., Benhamou, S., Börger,
L., Boyce, M.S., and Gaillard, J.-M. (2009) Memory keeps
you at home: a mechanistic model for home range emergence. Oikos, 118, 641–52.
van Noordwijk, A.J., Pulido, F., Helm, B., Coppack,
T., Delingat, J., Dingle, H., Hedenström, A., van der
Jeugd, H., Marchetti, C., Nilsson, A., and Pérez-Tris, J.
(2006) A framework for the study of genetic variation in
migratory behavior. Journal of Ornithology, 147, 221–33.
van Soest, P.J. (1982) Nutritional Ecology of the Ruminant,
O and B Books: Corvallis, OR.
Vanni, M.A., DeAngelis, D.L., Schindler, D.E., and Huxel,
G.R. (2004) Overview: cross-habitat flux of nutrients
and detritus. In G. E. Polis, M. E. Power, and G. R. Huxel
(eds) Food Webs at the Landscape Level, pp. 3–11,
University of Chicago Press: Chicago.
Varpe, Ø., Fiksen, Ø., and Slotte, A. (2005) Meta-ecosystems
and biological energy transport from ocean to coast: the
ecological importance of herring migration. Oecologia,
146, 443–51.
Veeranagoudar, D.K., Shanbhag, B.A., and Saidapur, S.K.
(2004) Foraging behaviour in tadpoles of the bronze
frog Rana temporalis: experimental evidence for the
Ideal Free Distribution. Journal of Biosciences, 29, 2,
201–207.
Ventsel, A. (2006) Hunting–herding continuum in Sakha.
In F. Stammler and H. Beach (eds) People and Reindeer
on the Move. Special Issue of the journal Nomadic
Peoples, 10, 2, 68–86, Berghahn Publishers: Oxford.
Videler, J. and Groenewold, A. (1991) Field measurements
of hanging flight aerodynamics in the kestrel Falco tinnurculus. Journal of Experimental Biology, 155, 519–30.
Visser M.E., Holleman, L.J.M., and Gienapp, P. (2006)
Shifts in caterpillar biomass phenology due to climate
change and its impact on the breeding biology of an
insectivorous bird. Oecologia, 147, 164–72.
Vitebsky, P. (2005) Reindeer People. Living with Animals
and Spirits in Siberia, Harper Collins: London.
Vogel, S. (1994) Life in Moving Fluids: The Physical
Biology of Flow, Princeton University Press: Princeton.
von Essen, L. (1991) A note on the Lesser White-fronted
goose. Ardea, 79, 305–306.
von Haartman, L. (1968) The evolution of resident versus
migratory habit in birds. Some considerations. Ornis
Fennica, 45, 1–6.
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Vors, L.S., Schaefer, J.A., Pond, B.A., Rodgers, A.R., and
Patterson, B.R. (2007) Woodland Caribou Extirpation
and Anthropogenic Landscape Disturbance in Ontario.
Journal of Wildlife Management, 71, 4, 1249–56.
Vrieze, L.A. and Sorensen, P.W. (2001) Laboratory
assessment of the role of a larval pheromone and natural stream odor in spawning stream localization by
migratory sea lamprey (Petromyzon murinus).
Canadian Journal of Fisheries and Aquatic Sciences,
58, 2374–85.
Wacher, T.J, Rawlings, P., and W.F. Snow. (1993) Cattle
migration and stocking densities in relation to tsetsetrypanosomiasis challenge in the Gambia. Annals of
Tropical Medicine and Parasitology, 87, 517–24.
Walker, M.M., Diebel, C.E., Haugh, C.V., Pankhurst, P.M.,
Montgomery, J.C., and Green, C.R. (1997) Structure and
function of the vertebrate magnetic sense. Nature, 390,
371–76.
Walker, N.A., Henry, H.A.L., Wilson, D.J., and Jefferies,
R.L. (2003) The dynamics of nitrogen movement in an
Arctic salt marsh in response to goose herbivory: a
parameterized model with alternate stable states.
Journal of Ecology, 91, 637–50.
Wall, J., Douglas-Hamilton, I., and Vollrath, F. (2006)
Elephants avoid costly mountaineering. Current
Biology, 16, R527–29.
Wallraff, H.G. (2004) Avian olfactory navigation: Its empirical foundation and conceptual state. Animal Behaviour,
67, 189–204.
Wallraff, H.G. and Andreae, M.O. (2000) Spatial gradients
in ratios of atmospheric trace gases: a study stimulated
by experiments on bird navigation. Tellus Series
B-Chemical And Physical Meteorology, 52, 1138–57.
Walther, G.-R., Post, E., Convey, P., Menzel, A., Parmesan,
C., Beebee, T.J.C., Fromentin, J.-M.,Hoegh-Guldberg, O.,
and Bairlein, F. (2002) Ecological responses to recent climate change. Nature, 416, 389–395.
Wang, W.L. (2003) China case study I: Studies on traditional transhumance and a system where herders return
to settled winter bases in Burjin County, Altai Prefecture,
Xinjiang, China. In J.M. Suttie and S.G. Reynolds (eds)
Transhumant grazing systems in temperate Asia, pp.
115–42, Food and Agriculture Organization of the
United Nations (FAO): Rome.
Ward, E.J. (2008) A review and comparison of four commonly used Bayesian and maximum likelihood model
selection tools. Ecological Modelling, 211, 1–10.
Ward, J.F., Austin R.M., and Macdonald, D.W. (2000) A
simulation model of foraging behaviour and the effect
of predation risk. Journal of Animal Ecology, 69, 16–30.
Ward, S., Bishop, C.M., Woakes, A.J., and Butler, P.J. (2002)
Heart rate and the rate of oxygen consumption of flying
and walking barnacle geese (Branta leucopsis) and
253
bar-headed geese (Anser indicus). Journal of Experimental
Biology, 205, 3347–56.
Watkinson, A. and Sutherland, W.J. (1995) Sources, sinks
and pseudo-sinks. Journal of Animal Ecology, 64, 126–30.
Watson, J.W., Epperly, S.P., Shah, A.K., and Foster, D.G.
(2005) Fishing methods to reduce sea turtle mortality
associated with pelagic longlines. Canadian Journal of
Fisheries and Aquatic Sciences, 62, 965–81.
Watson, M., Wilson, J.M., Koshkin, M., Sherbakov, B.,
Karpov, F., Gavrilov, A., Schielzeth, H., Brombacher, M.,
Collar, N.J., and Cresswell, W. (2006) Nest survival and
productivity of the critically endangered sociable lapwing, Vanellus gregarius. Ibis, 148, 489–502.
Weber, J.M. (2009) The physiology of long-distance migration: extending the limits of endurance metabolism.
Journal of Experimental Biology, 212, 593–97.
Weber, R.E., Jessen, T.H., Malte, H., and Tame, J. (1993)
Mutant hemoglobins (a119-ala and b55-ser) functions
related to high-altitude respiration in geese. Journal of
Applied Physiology, 75, 2646–55.
Weber, T.P. and Hedenström, A. (2001) Long-distance
migrants as a model system of structural and physiological plasticity. Evolutionary Ecology Research, 3,
255–71.
Weber, T.P. and Houston, A. I. (1997) Flight costs, flight
range and stopover ecology of migrating birds. Journal
of Animal Ecology, 66, 297–306.
Weber, T.P., Fransson, T., and Houston, A.I. (1999) Should
I stay or should I go? Testing optimality models of stopover decisions in migrating birds. Behavioral Ecology
and Sociobiology, 46, 280–86.
Webster, M.S., Marra, P.P., Haig, S.M., Bensch, S., and
Holmes, R.T. (2002) Links between worlds: unraveling
migratory connectivity. Trends in Ecology and Evolution,
17, 76–83.
Wedin, W.F., Lingvall, P., Thorsell, B, and Jonsson, N.
(1984) Quality of first-growth forage during maturation at diverse latitudes. In H. Riley and A.O Skjelvag
(eds) The Impact of Climate on Grass Production and
Quality, Proceedings of the 10th General Meeting of
the European Grassland Federation, As, Norway, pp.
46–50.
Weimerskirch, H. (2010) Editorial. Argos Forum, 68, 3.
http://argos-system.clsamerica.com/documents/publications/newsletter/anl_68.pdf.
Weimerskirch, H. and Wilson, R.P. (2000) Oceanic respite
for wandering albatrosses. Nature, 406, 955–56.
Weiner, D.R. (2000) Models of Nature: Ecology,
Conservation, and Cultural Revolution in Soviet
Russia, University of Pittsburgh Press: Pittsburgh,
324 pp.
Weng, K.C., Boustany, A.M., Pyle, P. Anderson, S.D.,
Brown, A., and Block, B.A. (2007) Migration and habitat
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
254
REFERENCES
of white sharks (Carcharodon carcharias) in the eastern
Pacific Ocean. Marine Biology, 152, 877–94.
Western, D. (1975) Seasonality in water availability and its
influence of the structure, dynamics and efficiency of a
savannah large mammal community. East African
Wildlife Journal, 13, 265–86.
White, P.J., Davis, T.L., Barnowe-Meyer, K.K., Crabtree,
R.L., and Garrott, R.A. (2007) Partial migration and
philopatry of Yellowstone pronghorn. Biological
Conservation, 135, 502–10.
White, R.G. (1983) Foraging patterns and their multiplier
effects on productivity of northern ungulates. Oikos, 40,
377–84.
Whyte, I.J. and Joubert, S.C.J. (1988) Blue wildebeest population trends in the Kruger National-Park and the
effects of fencing. South African Journal of Wildlife
Research, 18, 78–87.
Widmer, M. (1999) Altitudinal variation of migratory traits
in the garden warbler Sylvia borin. PhD thesis, University
of Zürich.
Wiens, J.A. (1989) The Ecology of Bird Communities. Volume
1. Foundations and Patterns. Cambridge Studies in
Ecology, Cambridge University Press: Cambridge.
Wiens, J.A. (1991) Ecological similarity of shrub-desert
avifaunas of Australia and North America. Ecology, 72,
479–95.
Wikelski, M. and Cooke S.J., (2006) Conservation physiology. Trends in Ecology and Evolution, 21, 38–46.
Wikelski, M., Kays, R.W., Kasdin, N.J., Thorup, K., Smith,
J.A., and Swenson, G.W., Jr. (2007) Going wild: what a
global small-animal tracking system could do for experimental biologists. Journal of Experimental Biology, 210,
181–86.
Wikelski, M., Moskowith, D., Adelman, J.S., Cochran,
J., Wilcove, D.S. and May, M.L. (2006) Simple rules guide
dragonfly migration. Biology Letters, 2, 325–29.
Wikelski, M., Tarlow, E., Raim, A., Diehl, R.H., Larkin,
R.P., and Visser, G.H. (2003) Costs of migration in freeflying songbirds. Nature, 423, 704.
Wilcove, D. (2007) No Way Home: The Decline of the World’s
Great Migrations. Island Press: Covello, CA, 256 pp.
Wilcove, D. S. (2008) Animal migration: An endangered
phenomenon? Issues in Science and Technology, 24,
71–78.
Wilcove, D.S. and Wikelski, M. (2008) Going, going gone:
Is animal migration disappearing. PLoS Biology, 6, e188.
doi:10.1371/journal.pbio.0060188.
Wilcove, D.S., Rothstein, D., Dubow, J., Phillips, A., and
Losos, E. (1998) Quantifying Threats to Imperiled
Species in the United States. Bioscience, 48, 8, 607–15.
Williams, R., Lusseau, D., and Hammond, P.S. (2009)
The role of social aggregations and protected areas
in killer whale conservation: the mixed blessing of critical
habitat. Biological Conservation, 142, 4, 709–19.
Williams, T.C., Ireland, L.C., and Williams, J.M. (1973) Highaltitude flights of free-tailed bat, Tadarida brasiliensis,
observed with radar. Journal of Mammalogy, 54, 807–21.
Williams, T.C. and Williams, J. M. (1978) The orientation of
transatlantic migrants. In K. Schmidt-Koenig and W.
Keeton (eds) Animal migration, orientation and homing, pp. 239–51, Springer-Verlag: Berlin.
Williamson, D., Williamson, J., and Ngwamotsoko, K.T.
(1988) Wildebeest migration in the Kalahari. African
Journal of Ecology, 26, 269–80.
Wilmshurst, J.F., Fryxell, J.M., and Hudson, R.J. (1995)
Forage quality and patch choice by wapiti (Cervus elaphus) Behavioral Ecology, 6, 2, 209–17.
Wilmshurst, J.F., Fryxell, J.M., and Bergman, C.M. (2000)
The allometry of patch selection in ruminanats. Proc.
Roy. Sco. Lond. (B), 267, 345–49.
Wilmshurst, J.F., Fryxell, J.M., Farm, B.P., Sinclair, A.R.E.,
and C.P. Henschel (1999) Spatial distribution of Serengeti
wildebeest in relation to resources. Canadian Journal of
Zoology, 77, 1223–32.
Wilps, H. and Diop, B. (1997) Field investigations on
Schistocerca gregaria (Forskal) adults, hoppers and hopper bands. In S. Krall, R. Peveling, and D. Ba Diallo (eds)
New Strategies in Locust Control., pp. 117–28,
Birkhäuser-Verlag: Basel.
Wilson, C. and Tisdell, C. (2003) Conservation and economic benefits of wildlife-based marine tourism: sea
turtles and whales as case studies. Human Dimensions
of Wildlife, 8, 1, 49–58.
Wilson, R. P., White, C.R., Quintana, F., Halsey, L.G.,
Liebsch, N., Martin, G.R., and Butler, P.J. (2006) Moving
towards acceleration for estimates of activity-specific
metabolic rate in free-living animals: the case of the cormorant. Journal of Animal Ecology, 75, 1081–90.
Wilson, S.G., Taylor, J.G., and Pearce, A.F. (2001) The seasonal
aggregation of whale sharks at Ningaloo Reef, Western
Australia: Currents, migrations and the El Nino/Southern
Oscillation. Environmental Biology of Fishes, 61, 1–11.
Wiltschko, R. and Wiltschko, W. (2003) Mechanisms of orientation and navigation in migratory birds. In
P. Berthold, E. Gwinner and E. Sonnenschein (eds) Avian
Migration, pp. 433–56, Springer Verlag: Berlin.
Wiltschko, W. and Wiltschko, R. (1972) Magnetic compass
of European robins. Science, 176, 62–64.
Wingfield, J.C. (2008) Organization of vertebrate annual
cycles: implications for control mechanisms. Philosophical
Transactions of the Royal Society of London, Series B,
363, 425–41.
Winkler, D.W. (2005) How do migration and dispersal
interact? In R. Greenberg and P.P. Marra (eds) Birds of
OUP CORRECTED PROOF – FINALS, 12/11/2010, SPi
R EFER EN C ES
Two Worlds: the Ecology and Evolution of Migration,
pp. 401–13, Johns Hopkins University Press: Baltimore.
Winkler, D.W. and Allen, P.E. (1996) The seasonal decline
in tree swallow clutch size: Physiological constraint or
strategic adjustment? Ecology, 77, 922–32.
Winkler, K. (2000) Migration and speciation. Nature,
404, 36.
Wirestam, R., Fagerlund, T., Rosén, M., and Hedenström,
A. (2008) Magnetic resonance imaging for non-invasive
analysis of fat storage in migratory. Auk, 125, 965–71.
Witey, J.C., Bloxton, T.D., and Marzluff, J.M. (2001) Effects
of tagging and location error in wildlife radiotelemetry
studies. In J. J. Millspaugh and J. M. Marzluff (eds)
Radiotracking and animal populations, pp. 45–75,
Academic Press: San Diego.
Woinarsky, J.C.Z. (2006) Predictors of nomadism in
Australian birds: a reanalysis of Allen and Saunders
(2002) Ecosystems. 9, 689–93.
Wolanski, E. and Gereta, E. (2001) Water quality and quantity as the factors driving the Serengeti ecosystem.
Tanzania. Hydrobiologia. 458, 169–80.
Wolf, C.M., Griffith, B., Reed, C., and Temple, S.A. (1996)
Avian and mammalian translocations: update and reanalysis of 1987 survey data. Conservation Biology, 10,
1142–54.
Wood, C.R., Chapman, J.W., Reynolds, D.R., Barlow, J.F.,
Smith, A.D., and Woiwod, I.P. (2006) The influence of the
atmospheric boundary layer on nocturnal layers of noctuids and other moths migrating over southern Britain.
International Journal of Biometeorology, 50, 193–204.
Wood, P. and Wolfe, M.L. (1988) Intercept feeding as a
means of reducing deer-vehicle collisions. Wildlife
Society Bulletin, 16, 376–80.
Woodbury, A.M. (1941) Animal migration: periodic-response theory. The Auk, 58, 463–505.
Woodrey, M.S. and Chandler, C.R. (1997) Age-related timing of migration: Geographic and interspecific patterns.
Wilson Bulletin, 109, 52–67.
Woodrey, M.S. and Moore, F.R. (1997) Age-related differences in the stopover of fall landbird migrants on the
coast of Alabama. Auk, 114, 695–707.
Woods, J. (1991) Ecology of a partially migratory elk population Ph.D. thesis, University of British Columbia:
Vancouver, BC.
Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C.,
Halpern, B.S., Jackson, J.B.C., Lotze, H.K., Micheli,
F., Palumbi, S.R., Sala, E., Selkoe, K.A., Stachowicz,
J.J., and Watson, R. (2006) Impacts of Biodiversity Loss on
Ocean Ecosystem Services. Science, 314, 3, 787–90.
Wright, S. (1931) Evolution in Mendelian populations.
Genetics, 16, 97–159.
255
Xia, L., Yang, Q., Li, Z., Wu, Y., and Feng, Z. (2007)
The effect of the Qinghai-Tibet railway on the migration of Tibetan antelope (Pantholops hodgsonii)
in Hoh-xil National Nature Reserve, China. Oryx, 41,
3, 352–57.
Yan Zhaoli, Ning, Wu, Dorji, Yeshi, and Jia, Ru (2005) A
review of rangeland privatisation and its implications
in the Tibetan Plateau, China. Nomadic Peoples, 9 (1 &
2), 31–52.
Yates, C.A., R. Erban, C. Escudero, I.D. Couzin, J. Buhl, I.
G. Kevrekidis, P.K. Maini, and D.J.T. Sumpter (2009)
Inherent noise can facilitate coherence in collective
swarm motion. Proceedings of the National Academy of
Sciences, 106, 5464–69.
Ydenberg, R.C., Butler, R.W., Lank, D.B., Guglielmo,
C.G., Lemon, M., and Wolf, N. (2002) Trade-offs,
condition dependence and stopover site selection
by migrating sandpipers. Journal of Avian Biology,
33, 47–55.
Yeh, E.T. (2005) Green governmentality and pastoralism in
Western China: ‘Converting pastures to grasslands’.
Nomadic Peoples, 9, 9–30.
Yong, W., Finch, D.M., Moore, F.R., and Kelly, J.F. (1998)
Stopover ecology and habitat use of migratory Wilson’s
warblers. Auk, 115, 829–42.
Yoshida, A. (1997) Kul’tura Pitania Gydanskikh Nentsev
(Interpretatsia i sotsial’naia adaptatsiia) Novye Issledovania
po etnologii i antropologii, Russian Academy of Sciences:
Moscow.
Yoshihara, Y., Ito, T.Y., Lhagvasuren, B., and Takatsuki, S.
(2008) A comparison of food resources used by
Mongolian gazelles and sympatric livestock in three
areas in Mongolia. Journal of Arid Environments, 72, 1,
48–55, DOI: 10.1016/j.jaridenv.2007.05.001.
Young, K.A. (1999) Managing the decline of Pacific salmon:
Metapopulation theory and artificial recolonization as
ecological mitigation. Canadian Journal of Fisheries and
Aquatic Sciences, 56, 9, 1700–706.
Yuzhakov, A. and Mukhachev, A. (2001) Etnicheskoe
Olenevodstvo Zapadnoi Sibiri: Nenetskii Tip, Agricultural Science Publishers: Novosibirsk.
Zabrodin, V.A., Borozdin, E.K., Vostriakov, P.N.,
D’iachenko, N.O., Kriuchkov, V.V., and Andreev, V.N.
(1979) Severnoe Olenevodstvo, Kolos: Moscow.
Zink, R.M. (2002) Towards a framework for understanding the evolution of avian migration. Journal of Avian
Biology, 33, 436.
Zwarts, L. and Dirksen, S. (1990) Digestive bottleneck limits the increase in food-intake of whimbrels preparing
for spring migration from the Banc-Darguin, Mauritania.
Ardea, 78, 257–78.
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
Animal Migration
A Synthesis
edited by
E.J. Milner-Gulland,
John M. Fryxell,
and
Anthony R.E. Sinclair
1
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
1
Great Clarendon Street, Oxford ox2 6dp
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ISBN 978–0–19–956899–4 (Hbk.)
978–0–19–956900–7 (Pbk.)
1 3 5 7 9 10 8 6 4 2
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
Contents
List of contributors
vii
Acknowledgements
ix
1 Introduction
John M. Fryxell, E.J. Milner-Gulland, and Anthony R.E. Sinclair
1
Part 1: The evolution of migration
5
2 Understanding the evolution of migration through empirical examples
Katherine A. Cresswell, William H. Satterthwaite, and Gregory A. Sword
7
3 Theoretical reflections on the evolution of migration
Robert D. Holt and John M. Fryxell
17
Part 2: How to migrate
33
4 Mechanistic principles of locomotion performance in migrating animals
Anders Hedenström, Melissa S. Bowlin, Ran Nathan, Bart A. Nolet, and Martin Wikelski
35
5 Energy gain and use during animal migration
Nir Sapir, Patrick J. Butler, Anders Hedenström, and Martin Wikelski
52
6 Cues and decision rules in animal migration
Silke Bauer, Bart A. Nolet, Jarl Giske, Jason W. Chapman, Susanne Åkesson,
Anders Hedenström, and John M. Fryxell
68
Part 3: Migration in time and space
89
7 Uncertainty and predictability: the niches of migrants and nomads
Niclas Jonzén, Endre Knudsen, Robert D. Holt, and Bernt-Erik Sæther
91
8 Migration quantified: constructing models and linking them with data
Luca Börger, Jason Matthiopoulos, Ricardo M. Holdo, Juan M. Morales,
Iain Couzin, and Edward McCauley
111
Part 4: Broader contexts
129
9 Migration impacts on communities and ecosystems: empirical evidence and theoretical insights
Ricardo M. Holdo, Robert D. Holt, Anthony R.E. Sinclair, Brendan J. Godley, and Simon Thirgood
131
v
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vi
CONTENTS
10 Pastoral migration: mobile systems of animal husbandry
Roy H. Behnke, Maria E. Fernandez-Gimenez, Matthew D. Turner, and Florian Stammler
144
11 Conservation and management of migratory species
Jennifer L. Shuter, Annette C. Broderick, David J. Agnew, Niclas Jonzén, Brendan J. Godley,
E.J. Milner-Gulland, and Simon Thirgood
172
12 Conclusions
E.J. Milner-Gulland, John M. Fryxell, and Anthony R.E. Sinclair
207
References
Index
216
257
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
Contributors
David J. Agnew, Department of Life Sciences, Imperial
College London, UK
Susanne Åkesson, Department of Biology, Ecology
Building, Lund University, SE-223 62 Lund, Sweden
Silke Bauer, Netherlands Institute of Ecology (NIOOKNAW), PO Box 1299, 3600 BG Maarssen, The
Netherlands and Swiss Ornithological Institute,
Luzernstrasse, 6204 Sempach, Switzerland
Roy H. Behnke, The Odessa Centre, Great Wolford, UK
Luca Börger, Department of Integrative Biology, University
of Guelph, Guelph, Ontario, Canada, N1G 2W1
Melissa S. Bowlin, Department of Biology, Ecology
Building, Lund University, SE-223 62 Lund, Sweden
Annette C. Broderick, Centre for Ecology and
Conservation, University of Exeter, Cornwall Campus,
Penryn, TR10 9EZ, UK
Patrick J. Butler, Centre for Ornithology, School of
Biosciences, The University of Birmingham, UK
Jason W. Chapman, Plant and Invertebrate Ecology,
Rothamsted Research, Harpenden, Herts AL5 2JQ, UK
Iain Couzin, Princeton University, Department of Ecology
& Evolutionary Biology, Princeton, NJ 08544, USA
Katherine A. Cresswell, Center for Stock Assessment
Research, Mail Stop E2, Jack Baskin School of
Engineering, University of California, Santa Cruz,
California, 95064, USA
Maria E. Fernandez-Gimenez, Department of Forest,
Rangeland and Watershed Stewardship, Colorado State
University, Fort Collins, CO, USA
John M. Fryxell, Department of Integrative Biology,
University of Guelph, Guelph, Ontario, Canada, N1G
2W1
Jarl Giske, University of Bergen, Department of Biology,
Postboks 7803, 5020 Bergen, Norway
Brendan J. Godley, Centre for Ecology and Conservation,
University of Exeter, Cornwall Campus, Penryn, TR10
9EZ, UK
Anders Hedenström, Department of Biology, Ecology
Building, Lund University, SE-223 62 Lund, Sweden
Ricardo M. Holdo, University of Missouri, Columbia, MO
65211, USA
Robert D. Holt, Department of Biology, University of
Florida, Gainesville, FL 32611, USA
Niclas Jonzén, Department of Biology, Ecology Building,
Lund University, SE-223 62 Lund, Sweden
Endre Knudsen, Centre for Ecological and Evolutionary
Synthesis (CEES), Department of Biology, University of
Oslo, PO Box 1066, Blindern, 0316 Oslo, Norway
Jason Matthiopoulos, Scottish Oceans Institute, School of
Biology, University of St Andrews, St Andrews, Fife,
KY16 8LB, Scotland, UK
Edward McCauley, National Center for Ecological
Analysis and Synthesis, Santa Barbara, CA 93101,
USA
E.J. Milner-Gulland, Imperial College London, Silwood
Park Campus, Buckhurst Road, Ascot, SL5 7PY, UK
Juan Manuel Morales, Laboratorio Ecotono, INIBIOMACONICET, Universidad Nacional del Comahue, Quintral
1250, 8400 Bariloche, Argentina
Ran Nathan, Department of Evolution, Systematics and
Ecology, Alexander Silberman Institute of Life Sciences,
The Hebrew University of Jerusalem, Edmund J. Safra
Campus at Givat Ram, 91904 Jerusalem, Israel
Bart A. Nolet, Netherlands Institute of Ecology (NIOOKNAW), PO Box 1299, 3600 BG Maarssen, The
Netherlands
Nir Sapir, Department of Evolution, Systematics and
Ecology, Alexander Silberman Institute of Life Sciences,
The Hebrew University of Jerusalem, Edmund J. Safra
Campus at Givat Ram, 91904 Jerusalem, Israel
Bernt-Erik Sæther, Centre for Conservation Biology,
Department of Biology, Norwegian University of
Science and Technology, NO-7491 Trondheim, Norway
William H. Satterthwaite, Center for Stock Assessment
Research, Mail Stop E2, Jack Baskin School of
Engineering, University of California, Santa Cruz,
California, 95064, USA
Jennifer L. Shuter, Department of Integrative Biology,
University of Guelph, Guelph, Ontario, Canada, N1G
2W1
Anthony R.E. Sinclair, Department of Zoology, University
of British Columbia, Vancouver, BC, Canada, V6T 1Z4
vii
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
viii
C O N T R I B U TO R S
Florian Stammler, Arctic Centre, University of Lapland,
PL 122, FIN 96101 Rovaniemi, Finland and Scott Polar
Research Institute, University of Cambridge, Lensfield
Road, Cambridge, CB2 1ER, UK
Gregory A. Sword, School of Biological Sciences, The
University of Sydney, Sydney, NSW, 2006 Australia
Simon Thirgood, Macaulay Land Use Research Institute,
Craigiebuckler, Aberdeen, UK
Matthew D. Turner , Department of Geography, 160
Science Hall, 550 Park Street, University of
Wisconsin, Madison, WI 53706, USA
Martin Wikelski, Department of Migration & Immunoecology, Max Planck Institute for Ornithology,
Germany and Chair of Ornithology, Konstanz
University, 78457 Konstanz, Germany
OUP CORRECTED PROOF – FINALS, 12/09/2010, SPi
Acknowledgements
We thank the UK Natural Environment Research
Council’s Centre for Population Biology for funding and hosting the workshop upon which this
book is based.
N.S. was supported by the US–Israel Binational
Science Foundation (BSF grant numbers 229/2002
and 124/2004), the Robert Szold Fund for Applied
Science, the Ring Foundation, and Rieger-JNF fellowships. P.J.B. was supported by BBSRC (grant
number BB/F015615/1 awarded to himself and Dr
C.M. Bishop). RDH and JF thank NSF and NSERC
for supporting their research in this general area for
a number of years. RDH thanks the University of
Florida Foundation for support. EJMG acknowledges the support of a Royal Society Wolfson
Research Merit award.
We thank Penny Hancock for valuable discussions and input during the preparation of the book,
and Tal Avgar and Cortland Griswold for thoughtful comments on Chapter 3. EJMG thanks Franck
Courchamp for hosting her in his lab during the
final stages of the book.
We thank Ian Sherman and Helen Eaton at OUP
for all their support and patience throughout.
ix